As ecosystems are defined by the network of interactions among organisms,

and between organisms and their environment,[4] they can be of any size but
usually encompass specific, limited spaces[5] (although some scientists say
that the entire planet is an ecosystem).[6]
Nebulous- unclear

An ecosystem is a community of living organisms (plants, animals and

microbes) in conjunction with the nonliving componentsof their environment
(things like air, water and mineral soil), interacting as a system.[2] These biotic
and abiotic components are regarded as linked together through nutrient
cycles and energy flows.[3] As ecosystems are defined by the network of
interactions among organisms, and between organisms and their
environment,[4] they can be of any size but usually encompass specific, limited
spaces[5] (although some scientists say that the entire planet is an ecosystem).
[6]

Energy, water, nitrogen and soil minerals are other essential abiotic
components of an ecosystem. The energy that flows through ecosystems is
obtained primarily from the sun. It generally enters the system
through photosynthesis, a process that also captures carbon from the
atmosphere. By feeding on plants and on one another, animals play an
important role in the movement of matter and energy through the system.
They also influence the quantity of plant and microbial biomass present. By
breaking down dead organic matter, decomposers release carbon back to the
atmosphere and facilitate nutrient cycling by converting nutrients stored in
dead biomass back to a form that can be readily used by plants and other
microbes.[7]
Ecosystems are controlled both by external and internal factors. External
factors such as climate, the parent material which forms the soil
and topography, control the overall structure of an ecosystem and the way
things work within it, but are not themselves influenced by the ecosystem.
[8]
Other external factors include time and potential biota. Ecosystems are
dynamic entitiesinvariably, they are subject to periodic disturbances and are

in the process of recovering from some past disturbance.[9] Ecosystems in

similar environments that are located in different parts of the world can have
very different characteristics simply because they contain different species.
[8]
The introduction of non-native species can cause substantial shifts in
ecosystem function. Internal factors not only control ecosystem processes but
are also controlled by them and are often subject to feedback loops.[8] While
the resource inputs are generally controlled by external processes like climate
and parent material, the availability of these resources within the ecosystem is
controlled by internal factors like decomposition, root competition or shading.
[8]
Other internal factors include disturbance, succession and the types of
species present. Although humans exist and operate within ecosystems, their
cumulative effects are large enough to influence external factors like climate.[8]
Biodiversity affects ecosystem function, as do the processes
of disturbance and succession. Ecosystems provide a variety ofgoods and
services upon which people depend; the principles of ecosystem
management suggest that rather than managing individual species, natural
resources should be managed at the level of the ecosystem itself. Classifying
ecosystems into ecologically homogeneous units is an important step towards
effective ecosystem management, but there is no single, agreed-upon way to
do this.
Contents
[hide]

1 History and development

2 Ecosystem processes
o

2.1 Primary production

2.2 Energy flow

2.3 Decomposition

2.4 Nutrient cycling

2.5 Function and biodiversity

2.6 Ecosystem goods and services

2.7 Ecosystem management

3 Ecosystem dynamics
o

3.1 Ecosystem ecology

4 Classification

4.1 Types

5 Anthropogenic threats

6 See also

7 Notes

8 References

9 Literature cited

10 External links

History and development[edit]

The term "ecosystem" was first used in a publication by British
ecologist Arthur Tansley.[fn 1][10] Tansley devised the concept to draw
attention to the importance of transfers of materials between organisms
and their environment.[11] He later refined the term, describing it as "The
whole system, ... including not only the organism-complex, but also the
whole complex of physical factors forming what we call the
environment".[12] Tansley regarded ecosystems not simply as natural
units, but as mental isolates.[12] Tansley later[13] defined the spatial extent
of ecosystems using the term ecotope.

G. Evelyn Hutchinson, a pioneering limnologist who was a contemporary of

Tansley's, combined Charles Elton's ideas about trophic ecology with those of
Russian geochemist Vladimir Vernadsky to suggest that mineral nutrient
availability in a lake limited algal production which would, in turn, limit the
abundance of animals that feed on algae. Raymond Lindeman took these
ideas one step further to suggest that the flow of energy through a lake was
the primary driver of the ecosystem. Hutchinson's students, brothers Howard
T. Odum and Eugene P. Odum, further developed a "systems approach" to the
study of ecosystems, allowing them to study the flow of energy and material
through ecological systems.[11]

Ecosystem processes[edit]
Energy and carbon enter ecosystems through photosynthesis, are
incorporated into living tissue, transferred to other organisms that feed on the
living and dead plant matter, and eventually released through respiration.
[14]
Most mineral nutrients, on the other hand, are recycled within ecosystems. [15]
Ecosystems are controlled both by external and internal factors. External
factors, also called state factors, control the overall structure of an ecosystem
and the way things work within it, but are not themselves influenced by the
ecosystem. The most important of these is climate.[8] Climate determines
the biome in which the ecosystem is embedded. Rainfall patterns and
temperature seasonality determine the amount of water available to the
ecosystem and the supply of energy available (by influencing photosynthesis).
[8]
Parent material, the underlying geological material that gives rise to soils,
determines the nature of the soils present, and influences the supply of
mineral nutrients. Topography also controls ecosystem processes by affecting
things like microclimate, soil development and the movement of water through
a system. This may be the difference between the ecosystem present
in wetland situated in a small depression on the landscape, and one present
on an adjacent steep hillside.[8]
Other external factors that play an important role in ecosystem functioning
include time and potential biota. Ecosystems are dynamic entitiesinvariably,

they are subject to periodic disturbances and are in the process of recovering
from some past disturbance.[9] Time plays a role in the development of soil
from bare rock and the recovery of a community from disturbance.[8] Similarly,
the set of organisms that can potentially be present in an area can also have a
major impact on ecosystems. Ecosystems in similar environments that are
located in different parts of the world can end up doing things very differently
simply because they have different pools of species present.[8]The introduction
of non-native species can cause substantial shifts in ecosystem function.
Unlike external factors, internal factors in ecosystems not only control
ecosystem processes, but are also controlled by them. Consequently, they are
often subject tofeedback loops.[8] While the resource inputs are generally
controlled by external processes like climate and parent material, the
availability of these resources within the ecosystem is controlled by internal
factors like decomposition, root competition or shading.[8] Other factors like
disturbance, succession or the types of species present are also internal
factors. Human activities are important in almost all ecosystems. Although
humans exist and operate within ecosystems, their cumulative effects are
large enough to influence external factors like climate.[8]

Primary production[edit]

Global oceanic and terrestrial phototroph abundance, from September 1997 to

August 2000. As an estimate of autotrophbiomass, it is only a rough indicator of
primary production potential, and not an actual estimate of it. Provided by
the SeaWiFS Project,NASA/Goddard Space Flight Center and ORBIMAGE.

Main article: Primary production

Primary production is the production of organic matter from inorganic carbon
sources. Overwhelmingly, this occurs through photosynthesis. The energy
incorporated through this process supports life on earth, while the carbon
makes up much of the organic matter in living and dead biomass, soil
carbon and fossil fuels. It also drives the carbon cycle, which influences
global climate via the greenhouse effect.
Through the process of photosynthesis, plants capture energy from light and
use it to combinecarbon dioxide and water to
produce carbohydrates and oxygen. The photosynthesis carried out by all the
plants in an ecosystem is called the gross primary production (GPP).[16] About
4860% of the GPP is consumed in plant respiration. The remainder, that
portion of GPP that is not used up by respiration, is known as the net primary
production (NPP).[14] Total photosynthesis is limited by a range of
environmental factors. These include the amount of light available, the amount
of leaf area a plant has to capture light (shading by other plants is a major
limitation of photosynthesis), rate at which carbon dioxide can be supplied to
the chloroplasts to support photosynthesis, the availability of water, and the
availability of suitable temperatures for carrying out photosynthesis.[16]

Energy flow[edit]

Left: Energy flow diagram of a frog. The frog represents a node in an extended food
web. The energy ingested is utilized for metabolic processes and transformed into
biomass. The energy flow continues on its path if the frog is ingested by predators,
parasites, or as a decaying carcass in soil. This energy flow diagram illustrates how
energy is lost as it fuels the metabolic process that transforms the energy and

nutrients into biomass.

Right: An expanded three link energy food chain (1. plants, 2. herbivores, 3.
carnivores) illustrating the relationship between food flow diagrams and energy
transformity. The transformity of energy becomes degraded, dispersed, and
diminished from higher quality to lesser quantity as the energy within a food chain
flows from one trophic species into another. Abbreviations: I=input, A=assimilation,
R=respiration, NU=not utilized, P=production, B=biomass. [17]

Main article: Energy flow (ecology)

See also: Food web and Trophic level
The carbon and energy incorporated into plant tissues (net primary
production) is either consumed by animals while the plant is alive, or it
remains uneaten when the plant tissue dies and becomesdetritus. In terrestrial
ecosystems, roughly 90% of the NPP ends up being broken down
bydecomposers. The remainder is either consumed by animals while still alive
and enters the plant-based trophic system, or it is consumed after it has died,
and enters the detritus-based trophic system. In aquatic systems, the
proportion of plant biomass that gets consumed by herbivores is much higher.
[18]
In trophic systems photosynthetic organisms are the primary producers. The
organisms that consume their tissues are called primary consumers
or secondary producersherbivores. Organisms which feed
on microbes (bacteria and fungi) are termed microbivores. Animals that feed
on primary consumerscarnivoresare secondary consumers. Each of
these constitutes a trophic level.[18] The sequence of consumptionfrom plant
to herbivore, to carnivoreforms a food chain. Real systems are much more
complex than thisorganisms will generally feed on more than one form of
food, and may feed at more than one trophic level. Carnivores may capture
some prey which are part of a plant-based trophic system and others that are
part of a detritus-based trophic system (a bird that feeds both on herbivorous
grasshoppers and earthworms, which consume detritus). Real systems, with
all these complexities, form food websrather than food chains.[18]

Decomposition[edit]
See also: Decomposition

The carbon and nutrients in dead organic matter are broken down by a group
of processes known as decomposition. This releases nutrients that can then
be re-used for plant and microbial production, and returns carbon dioxide to
the atmosphere (or water) where it can be used for photosynthesis. In the
absence of decomposition, dead organic matter would accumulate in an
ecosystem and nutrients and atmospheric carbon dioxide would be depleted.
[19]
Approximately 90% of terrestrial NPP goes directly from plant to
decomposer.[18]
Decomposition processes can be separated into three categoriesleaching,
fragmentation and chemical alteration of dead material. As water moves
through dead organic matter, it dissolves and carries with it the water-soluble
components. These are then taken up by organisms in the soil, react with
mineral soil, or are transported beyond the confines of the ecosystem (and are
considered "lost" to it).[19] Newly shed leaves and newly dead animals have
high concentrations of water-soluble components, and include sugars, amino
acids and mineral nutrients. Leaching is more important in wet environments,
and much less important in dry ones.[19]
Fragmentation processes break organic material into smaller pieces, exposing
new surfaces for colonization by microbes. Freshly shed leaf litter may be
inaccessible due to an outer layer of cuticle or bark, and cell contents are
protected by a cell wall. Newly dead animals may be covered by
an exoskeleton. Fragmentation processes, which break through these
protective layers, accelerate the rate of microbial decomposition.[19] Animals
fragment detritus as they hunt for food, as does passage through the
gut.Freeze-thaw cycles and cycles of wetting and drying also fragment dead
material.[19]
The chemical alteration of dead organic matter is primarily achieved through
bacterial and fungal action. Fungal hyphae produce enzymes which can break
through the tough outer structures surrounding dead plant material. They also
produce enzymes which break down lignin, which allows to them access to
both cell contents and to the nitrogen in the lignin. Fungi can transfer carbon

and nitrogen through their hyphal networks and thus, unlike bacteria, are not
dependent solely on locally available resources.[19]
Decomposition rates vary among ecosystems. The rate of decomposition is
governed by three sets of factorsthe physical environment (temperature,
moisture and soil properties), the quantity and quality of the dead material
available to decomposers, and the nature of the microbial community itself.
[20]
Temperature controls the rate of microbial respiration; the higher the
temperature, the faster microbial decomposition occurs. It also affects soil
moisture, which slows microbial growth and reduces leaching. Freeze-thaw
cycles also affect decompositionfreezing temperatures kill soil
microorganisms, which allows leaching to play a more important role in
moving nutrients around. This can be especially important as the soil thaws in
the Spring, creating a pulse of nutrients which become available.[20]
Decomposition rates are low under very wet or very dry conditions.
Decomposition rates are highest in wet, moist conditions with adequate levels
of oxygen. Wet soils tend to become deficient in oxygen (this is especially true
in wetlands), which slows microbial growth. In dry soils, decomposition slows
as well, but bacteria continue to grow (albeit at a slower rate) even after soils
become too dry to support plant growth. When the rains return and soils
become wet, the osmotic gradient between the bacterial cells and the soil
water causes the cells to gain water quickly. Under these conditions, many
bacterial cells burst, releasing a pulse of nutrients.[20]Decomposition rates also
tend to be slower in acidic soils.[20] Soils which are rich in clay minerals tend to
have lower decomposition rates, and thus, higher levels of organic matter.
[20]
The smaller particles of clay result in a larger surface area that can hold
water. The higher the water content of a soil, the lower the oxygen
content[21]and consequently, the lower the rate of decomposition. Clay minerals
also bind particles of organic material to their surface, making them less
accessibly to microbes.[20]Soil disturbance like tilling increase decomposition by
increasing the amount of oxygen in the soil and by exposing new organic
matter to soil microbes.[20]

The quality and quantity of the material available to decomposers is another

major factor that influences the rate of decomposition. Substances like sugars
and amino acids decompose readily and are considered
"labile". Cellulose and hemicellulose, which are broken down more slowly, are
"moderately labile". Compounds which are more resistant to decay, like lignin
or cutin, are considered "recalcitrant".[20] Litter with a higher proportion of labile
compounds decomposes much more rapidly than does litter with a higher
proportion of recalcitrant material. Consequently, dead animals decompose
more rapidly than dead leaves, which themselves decompose more rapidly
than fallen branches.[20] As organic material in the soil ages, its quality
decreases. The more labile compounds decompose quickly, leaving an
increasing proportion of recalcitrant material. Microbial cell walls also contain
recalcitrant materials like chitin, and these also accumulate as the microbes
die, further reducing the quality of oldersoil organic matter.[20]

Nutrient cycling[edit]
See also: Nutrient cycle, Biogeochemical cycle and Nitrogen cycle

Biological nitrogen cycling

Ecosystems continually exchange energy and carbon with the

wider environment; mineral nutrients, on the other hand, are mostly cycled
back and forth between plants, animals, microbes and the soil. Most nitrogen

enters ecosystems through biological nitrogen fixation, is deposited through

precipitation, dust, gases or is applied as fertilizer.[15] Since most terrestrial
ecosystems are nitrogen-limited, nitrogen cycling is an important control on
ecosystem production.[15]
Until modern times, nitrogen fixation was the major source of nitrogen for
ecosystems. Nitrogen fixing bacteria either live symbiotically with plants, or
live freely in the soil. The energetic cost is high for plants which support
nitrogen-fixing symbiontsas much as 25% of GPP when measured in
controlled conditions. Many members of the legume plant family support
nitrogen-fixing symbionts. Some cyanobacteria are also capable of nitrogen
fixation. These arephototrophs, which carry out photosynthesis. Like other
nitrogen-fixing bacteria, they can either be free-living or have symbiotic
relationships with plants.[15] Other sources of nitrogen includeacid
deposition produced through the combustion of fossil fuels, ammonia gas
which evaporates from agricultural fields which have had fertilizers applied to
them, and dust.[15] Anthropogenic nitrogen inputs account for about 80% of all
nitrogen fluxes in ecosystems.[15]
When plant tissues are shed or are eaten, the nitrogen in those tissues
becomes available to animals and microbes. Microbial decomposition
releases nitrogen compounds from dead organic matter in the soil, where
plants, fungi and bacteria compete for it. Some soil bacteria use organic
nitrogen-containing compounds as a source of carbon, and
releaseammonium ions into the soil. This process is known as nitrogen
mineralization. Others convert ammonium to nitrite and nitrate ions, a process
known as nitrification. Nitric oxide and nitrous oxide are also produced during
nitrification.[15] Under nitrogen-rich and oxygen-poor conditions, nitrates and
nitrites are converted to nitrogen gas, a process known as denitrification.[15]
Other important nutrients
include phosphorus, sulfur, calcium, potassium, magnesium and manganese.
[22]
Phosphorus enters ecosystems through weathering. As ecosystems age
this supply diminishes, making phosphorus-limitation more common in older

landscapes (especially in the tropics).[22] Calcium and sulfur are also produced
by weathering, but acid deposition is an important source of sulfur in many
ecosystems. Although magnesium and manganese are produced by
weathering, exchanges between soil organic matter and living cells account
for a significant portion of ecosystem fluxes. Potassium is primarily cycled
between living cells and soil organic matter.[22]

Function and biodiversity[edit]

See also: Biodiversity and Ecosystem engineer

Loch Lomond in Scotland forms a relatively isolated ecosystem. The fish community
of this lake has remained stable over a long period until a number of introductions in
the 1970s restructured its food web.[23]

Spiny forest at Ifaty, Madagascar, featuring various Adansonia(baobab)

species, Alluaudia procera(Madagascar ocotillo) and other vegetation.

Ecosystem processes are broad generalizations that actually take place

through the actions of individual organisms. The nature of the organismsthe
species, functional groups and trophic levels to which they belongdictates
the sorts of actions these individuals are capable of carrying out, and the
relative efficiency with which they do so. Thus, ecosystem processes are
driven by the number of species in an ecosystem, the exact nature of each

individual species, and the relative abundance organisms within these

species.[24] Biodiversity plays an important role in ecosystem functioning.[25]
Ecological theory suggests that in order to coexist, species must have some
level of limiting similaritythey must be different from one another in some
fundamental way, otherwise one species would competitively exclude the
other.[26] Despite this, the cumulative effect of additional species in an
ecosystem is not linearadditional species may enhance nitrogen retention,
for example, but beyond some level of species richness, additional species
may have little additive effect.[24] The addition (or loss) of species which are
ecologically similar to those already present in an ecosystem tends to only
have a small effect on ecosystem function. Ecologically distinct species, on
the other hand, have a much larger effect. Similarly, dominant species have a
large impact on ecosystem function, while rare species tend to have a small
effect. Keystone species tend to have an effect on ecosystem function that is
disproportionate to their abundance in an ecosystem.[24]

Ecosystem goods and services[edit]

Main articles: Ecosystem services and Ecological goods and services
See also: Ecosystem valuation and Ecological yield
Ecosystems provide a variety of goods and services upon which people
depend.[27] Ecosystem goods include the "tangible, material products"[28] of
ecosystem processesfood, construction material, medicinal plantsin
addition to less tangible items like tourism and recreation, and genes from wild
plants and animals that can be used to improve domestic species.
[27]
Ecosystem services, on the other hand, are generally "improvements in the
condition or location of things of value".[28] These include things like the
maintenance of hydrological cycles, cleaning air and water, the maintenance
of oxygen in the atmosphere, crop pollination and even things like beauty,
inspiration and opportunities for research.[27] While ecosystem goods have
traditionally been recognized as being the basis for things of economic value,
ecosystem services tend to be taken for granted.[28] While Gretchen Daily's
original definition distinguished between ecosystem goods and ecosystem

services, Robert Costanza and colleagues' later work and that of

the Millennium Ecosystem Assessment lumped all of these together as
ecosystem services.[28]

Ecosystem management[edit]
Main article: Ecosystem management
See also: Ecological economics, Sustainability and Sustainable development
When natural resource management is applied to whole ecosystems, rather
than single species, it is termed ecosystem management.[29] A variety of
definitions exist: F. Stuart Chapin and coauthors define it as "the application of
ecological science to resource management to promote long-term
sustainability of ecosystems and the delivery of essential ecosystem goods
and services",[30] while Norman Christensen and coauthors defined it as
"management driven by explicit goals, executed by policies, protocols, and
practices, and made adaptable by monitoring and research based on our best
understanding of the ecological interactions and processes necessary to
sustain ecosystem structure and function"[27] and Peter Brussard and
colleagues defined it as "managing areas at various scales in such a way that
ecosystem services and biological resources are preserved while appropriate
human use and options for livelihood are sustained".[31]
Although definitions of ecosystem management abound, there is a common
set of principles which underlie these definitions.[30] A fundamental principle is
the long-term sustainability of the production of goods and services by the
ecosystem;[30] "intergenerational sustainability [is] a precondition for
management, not an afterthought".[27] It also requires clear goals with respect
to future trajectories and behaviors of the system being managed. Other
important requirements include a sound ecological understanding of the
system, including connectedness, ecological dynamics and the context in
which the system is embedded. Other important principles include an
understanding of the role of humans as components of the ecosystems and
the use of adaptive management.[27] While ecosystem management can be
used as part of a plan for wilderness conservation, it can also be used in

intensively managed ecosystems[27] (see, for

example, agroecosystem and close to nature forestry).

Ecosystem dynamics[edit]

Temperate rainforest on theOlympic Peninsula in Washington state.

The High Peaks Wilderness Areain the 6,000,000-acre (2,400,000 ha)Adirondack

Park is an example of a diverse ecosystem.

Ecosystems are dynamic entitiesinvariably, they are subject to periodic

disturbances and are in the process of recovering from some past
disturbance.[9] When an ecosystem is subject to some sort of perturbation, it
responds by moving away from its initial state. The tendency of a system to
remain close to its equilibrium state, despite that disturbance, is termed
itsresistance. On the other hand, the speed with which it returns to its initial
state after disturbance is called its resilience.[9]

From one year to another, ecosystems experience variation in their biotic and
abiotic environments. A drought, an especially cold winter and a pest outbreak
all constitute short-term variability in environmental conditions. Animal
populations vary from year to year, building up during resource-rich periods
and crashing as they overshoot their food supply. These changes play out in
changes in NPP, decomposition rates, and other ecosystem processes.
[9]
Longer-term changes also shape ecosystem processesthe forests of
eastern North America still show legacies of cultivation which ceased 200
years ago, while methane production in eastern Siberian lakes is controlled by
organic matter which accumulated during the Pleistocene.[9]
Disturbance also plays an important role in ecological processes. F. Stuart
Chapin and coauthors define disturbance as "a relatively discrete event in
time and space that alters the structure of populations, communities and
ecosystems and causes changes in resources availability or the physical
environment".[32] This can range from tree falls and insect outbreaks to
hurricanes and wildfires to volcanic eruptions and can cause large changes in
plant, animal and microbe populations, as well soil organic matter content.
[9]
Disturbance is followed by succession, a "directional change in ecosystem
structure and functioning resulting from biotically driven changes in resources
supply."[32]
The frequency and severity of disturbance determines the way it impacts
ecosystem function. Major disturbance like a volcanic eruption or glacial
advance and retreat leave behind soils that lack plants, animals or organic
matter. Ecosystems that experience disturbances that undergo primary
succession. Less severe disturbance like forest fires, hurricanes or cultivation
result in secondary succession.[9] More severe disturbance and more frequent
disturbance result in longer recovery times. Ecosystems recover more quickly
from less severe disturbance events.[9]
The early stages of primary succession are dominated by species with small
propagules (seed and spores) which can be dispersed long distances. The
early colonizersoften algae, cyanobacteria and lichensstabilize the

substrate. Nitrogen supplies are limited in new soils, and nitrogen-fixing

species tend to play an important role early in primary succession. Unlike in
primary succession, the species that dominate secondary succession, are
usually present from the start of the process, often in the soil seed bank. In
some systems the successional pathways are fairly consistent, and thus, are
easy to predict. In others, there are many possible pathwaysfor example,
the introduced nitrogen-fixing legume, Myrica faya, alter successional
trajectories in Hawaiian forests.[9]
The theoretical ecologist Robert Ulanowicz has used information theory tools
to describe the structure of ecosystems, emphasizing mutual information
(correlations) in studied systems. Drawing on this methodology and prior
observations of complex ecosystems, Ulanowicz depicts approaches to
determining the stress levels on ecosystems and predicting system reactions
to defined types of alteration in their settings (such as increased or reduced
energy flow, and eutrophication.[33]

Ecosystem ecology[edit]
Main article: Ecosystem ecology
See also: Ecosystem model

A hydrothermal vent is an ecosystem on the ocean floor. (The scale bar is 1 m.)

Ecosystem ecology studies "the flow of energy and materials through

organisms and the physical environment". It seeks to understand the
processes which govern the stocks of material and energy in ecosystems, and
the flow of matter and energy through them. The study of ecosystems can
cover 10 orders of magnitude, from the surface layers of rocks to the surface
of the planet.[34]

There is no single definition of what constitutes an ecosystem.[35] German

ecologist Ernst-Detlef Schulze and coauthors defined an ecosystem as an
area which is "uniform regarding the biological turnover, and contains all the
fluxes above and below the ground area under consideration." They explicitly
reject Gene Likens' use of entire river catchments as "too wide a demarcation"
to be a single ecosystem, given the level of heterogeneity within such an area.
[36]
Other authors have suggested that an ecosystem can encompass a much
larger area, even the whole planet.[6] Schulze and coauthors also rejected the
idea that a single rotting log could be studied as an ecosystem because the
size of the flows between the log and its surroundings are too large, relative to
the proportion cycles within the log.[36] Philosopher of science Mark
Sagoff considers the failure to define "the kind of object it studies" to be an
obstacle to the development of theory in ecosystem ecology.[35]
Ecosystems can be studied through a variety of approachestheoretical
studies, studies monitoring specific ecosystems over long periods of time,
those that look at differences between ecosystems to elucidate how they work
and direct manipulative experimentation.[37] Studies can be carried out at a
variety of scales, frommicrocosms and mesocosms which serve as simplified
representations of ecosystems, through whole-ecosystem studies.[38] American
ecologist Stephen R. Carpenterhas argued that microcosm experiments can
be "irrelevant and diversionary" if they are not carried out in conjunction with
field studies carried out at the ecosystem scale, because microcosm
experiments often fail to accurately predict ecosystem-level dynamics.[39]
The Hubbard Brook Ecosystem Study, established in the White Mountains,
New Hampshire in 1963, was the first successful attempt to study an entire
watershed as an ecosystem. The study used stream chemistry as a means of
monitoring ecosystem properties, and developed a detailed biogeochemical
model of the ecosystem.[40]Long-term research at the site led to the discovery
of acid rain in North America in 1972, and was able to document the
consequent depletion of soil cations (especially calcium) over the next several
decades.[41]

Classification[edit]
See also: Ecosystem diversity, Ecoregion, Ecological land
classification and Ecotope

Flora of Baja California Desert,Catavia region, Mexico.

Classifying ecosystems into ecologically homogeneous units is an important

step towards effective ecosystem management.[42] A variety of systems exist,
based on vegetation cover, remote sensing, and bioclimatic
classificationsystems.[42] American geographer Robert Bailey defines a
hierarchy of ecosystem units ranging from microecosystems (individual
homogeneous sites, on the order of 10 square kilometres (4 sq mi) in area),
through mesoecosystems (landscape mosaics, on the order of 1,000 square
kilometres (400 sq mi)) to macroecosystems (ecoregions, on the order of
100,000 square kilometres (40,000 sq mi)).[43]
Bailey outlined five different methods for identifying ecosystems: gestalt ("a
whole that is not derived through considerable of its parts"), in which regions
are recognized and boundaries drawn intuitively; a map overlay system where
different layers likegeology, landforms and soil types are overlain to identify
ecosystems; multivariate clustering of site attributes; digital image
processing of remotely sensed data grouping areas based on their
appearance or other spectral properties; or by a "controlling factors method"
where a subset of factors (like soils, climate, vegetation physiognomy or
the distribution of plant or animal species) are selected from a large array of
possible ones are used to delineate ecosystems.[44] In contrast with Bailey's
methodology, Puerto Rico ecologist Ariel Lugo and coauthors identified ten

characteristics of an effective classification system: that it be based

on georeferenced, quantitative data; that it should minimize subjectivity and
explicitly identify criteria and assumptions; that it should be structured around
the factors that drive ecosystem processes; that it should reflect the
hierarchical nature of ecosystems; that it should be flexible enough to conform
to the various scales at which ecosystem management operates; that it should
be tied to reliable measures of climate so that it can "anticipat[e] global climate
change; that it be applicable worldwide; that it should be validated against
independent data; that it take into account the sometimes complex
relationship between climate, vegetation and ecosystem functioning; and that
it should be able to adapt and improve as new data become available".[42]

Types[edit]

Aquatic ecosystem

Marine ecosystem

Large marine ecosystem

Freshwater ecosystem

Lake ecosystem

River ecosystem

Wetland

Terrestrial ecosystem

Forest

Littoral zone

Riparian zone

Subsurface lithoautotrophic microbial ecosystem

Urban ecosystem

Desert

A freshwater ecosystem in Gran Canaria, an island of the Canary Islands.

Anthropogenic threats[edit]
See also: Planetary boundaries
As human populations grow, so do the resource demands imposed on
ecosystems and the impacts of the human ecological footprint. Natural
resources are not invulnerable and infinitely available. The environmental
impacts of anthropogenic actions, which are processes or materials derived
from human activities, are becoming more apparentair and water quality are
increasingly compromised, oceans are being overfished, pests and diseases
are extending beyond their historical boundaries, and deforestation is
exacerbating flooding downstream. It has been reported that approximately
4050% of Earth's ice-free land surface has been heavily transformed or
degraded by anthropogenic activities, 66% of marine fisheries are
either overexploited or at their limit, atmospheric CO2 has increased more than
30% since the advent ofindustrialization, and nearly 25% of Earth's bird
species have gone extinct in the last two thousand years.[45] Society is
increasingly becoming aware that ecosystem services are not only limited, but
also that they are threatened by human activities. The need to better consider
long-term ecosystem health and its role in enabling human habitation and
economic activity is urgent. To help inform decision-makers, many ecosystem

services are being assigned economic values, often based on the cost of
replacement with anthropogenic alternatives. The ongoing challenge of
prescribing economic value to nature, for example through biodiversity
banking, is prompting transdisciplinary shifts in how we recognize and
manage the environment, social responsibility, business opportunities, and our
future as a species.

Introduction
The following section describes some of the basic ecological concepts that
underlie the ACE Basin ecosystem. Descriptions include an introduction to
ecology and ecosystems, habitats, succession and biodiversity, populations and
communities, energy flow through ecosystems, and ecosystem services. Many
of these concepts are applicable to the habitats, communities, and ecosystems
that are described in the Biological Resources Section and other sections of this
product.
What is Ecology?
Ecology is the study of living organisms and their interactions with the physical
and biological environment. While this involves an incredibly complex network
of species, habitats, climates, physical environments, and human uses and
concerns, the field of ecology has continued to advance the explanation of
biological distributions, chemical cycles, and the interlinked nature of
ecosystems. Some of these biological distributions and ecosystem processes
(e.g. fish and decapod communities, carbon and nitrogen cycling) are described
in other sections of this characterization.
What is an Ecosystem?
The ACE Basin contains a diversity of habitats that range from subtidal areas

and vast wetlands to uplands. These habitats are populated by many different
plant and animal species that interact with the physical environment to create
the ACE Basin ecosystem. Ecosystems are defined as a set of organisms
(community) living in an area, their physical environment, and the interactions
between them (Daily 1997). For example, the ecology of Edisto Beach
consists of organisms such as fish, insects, shellfish, birds, raccoons, and
humans that make up the community; natural features such as the surf zone,
front beach, dunes, forested areas and the created infrastructure of roads,
buildings, and utilities that constitute the physical environment; and the
interactions between the community and physical components.
Although it has not always been clearly recognized, we are completely
dependent on the ecosystems in which we live. The myriad processes that
integrate energy and nutrients flowing through the ACE Basin ecosystem
provide its human inhabitants a variety of services. Besides providing food and
shelter, the ecosystem provides waste treatment (by way of carbon dioxide
consumption; oxygen production; and breakdown of sewage), a water filtration
system (by the soil), recreational opportunities, and a basis for economic
development. Remove any one of these services and the character and
function of the other components can be compromised. See the discussion of
ecosystem services below.
Natural geomorphic, as well as episodic, events have shaped the overall
diversity of habitats and the services they provide. Over the last 10,000 years as
early human societies created shell middens and set fires intentionally to
capture game, and especially over the last 200 years with the advent of
extensive forestry and agricultural practices, ecosystems have been
significantly altered by anthropogenic factors. These factors will likely
continue to pressure ecosystems in the future. (See related section: History.)
As habitats are modified, ecological processes in these habitats also change and
some of these changes may be significant. For example, estuarine marshes are
effective traps for many pollutants. When pollutants are introduced at low
levels, they may be bound to sediments or degraded through natural processes,
and they are less likely to be transferred up the food chain. If the ability of
sediments to trap such pollutants is exceeded, however as has happened in
some places in the Charleston Harbor Estuary (DHEC 1998), then pollutants
may move up the food chain. Some of the changes may not be immediately
obvious, such as a reduction in populations of benthic infauna (small worms,
crustaceans, and bivalves). Other impacts, such as the loss of oyster beds
resulting in the reduced availability of oysters for recreational or commercial
harvest, are much more obvious. Part of the solution to these problems lies in

appropriate management decisions made at the federal, state, and local

governmental levels, as well as by individual property owners and residents of
the ACE Basin. (See related section: Management.)
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Habitat
o Habitat Fragmentation
o Ecotonal Habitat
A habitat is defined in relation to a plant or animal species and is the location
where the species lives. It is a combination of the physical and biological
components of the location and ranges from very large and stable (the earth or
the open ocean) to very small and ephemeral (a pond in the dunes). A habitat
may be a stand of loblolly pine trees with heart-rot where red-cockaded
woodpeckers are found, the estuarine pluff mud where mud snails
and polychaete worms thrive, or the town of Edisto Beach where humans
reside. Usually, the more diverse the habitat types within a region, the greater
will be the variety of species being supported. Coastal areas, such as the ACE
Basin, located between the open ocean and upland areas, have a high diversity
of habitats and microhabitats, supporting diverse and abundant communities of
plants and animals. However, one of the greatest threats to habitat diversity in
the ACE Basin is the conversion of existing habitats to structurally and
biologically simpler habitats such as agricultural fields, pine plantations, and
urban or residential areas. In addition to the direct loss of existing habitat, the
resulting fragmentation of the remaining forested and wetland areas results in
decreased species diversity (Odum 1997; Meffe and Carroll 1994).
Habitat Fragmentation
Habitats are always patchy and fragmented, to some extent, by natural
disturbances and subsequent succession (see text below). While some of these
processes can have global impacts, such as climate change and sea level rise,
most disturbances occur on a more local scale. Local examples may include
forest fires, hurricanes, and disease. Through successional processes, these
habitats revert back to pre-impact states over time periods ranging from years
to decades. Low-level patches of disturbance can increase the variety of
habitats and therefore provide additional habitat for opportunistic species.

In contrast, due
to anthropogenic influences much of
the change has become much more
long-lasting. Forests have been
converted to agricultural fields and
suburban and urban land, and
seldom have the opportunity to
revert back to a natural state.
Depending on the land use,
anthropogenically altered habitats
tend to have a simpler structure with
lower habitat diversity and increased
edge habitats than do natural
systems. As a result, biotic
communities shift to favor those
species that can utilize open, edge,
agricultural, or suburban habitats.
Species that require interior forest habitats have a harder time finding
appropriate food and other required resources. Thus, there has been an overall
reduction in populations dependent on these habitats (Meffe and Carroll 1994).
Ecotonal Habitats
An ecotone is an area of transition between two different environments or
habitats. Some ecotones are extreme, such as the shift from
an estuarine or riverine environment to a forested habitat . In other places, the
transition may take place more gradually, shifting from open rivers to forested
wetlands and upland areas. One of the most obvious characteristics of an
ecotone is the shift in vegetation that occurs between adjacent habitats. These
changes reflect similar gradients in less obvious properties such as salinity,
moisture, flooding regime, altitude, and other physical and chemical properties.
Ecotonal or edge habitats often have a higher diversity of plants and animals
relative to the more homogenous habitats on either side. In addition to edgeselected species, species native to each adjacent habitat may occasionally be
found in these transition habitats.

One result of fragmented habitats arising

from development in the ACE Basin is the
change of the character of ecotonal habitats.
Ecotones where the vegetative communities
previously shifted slowly from wetland to
upland forest have been changed to sharper
boundaries between wetland areas and what
are now agricultural fields or suburban
developments. Many game species utilize
these edge habitats and a common practice
of wildlife managers is to create edgehabitat. However, many non-game species
require less abrupt transition areas and have
declined as a result of the loss of these
gradual transition zones (Meffe and Carroll
1994).
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Succession
Succession is a process that encompasses the dynamic biological and physical
changes that lead to the development of complex, interlinked, biological
communities. For example, habitats that are disturbed by fire or storm go
through a series of community changes as the habitat recovers from the
disturbance. Succession has been viewed as a step-wise process, whereby early
communities typically are dominated by a few opportunistic species. The
transfer of energy from primary producers (plants) to herbivores to predators
(each is considered a trophic level, see discussion of energy flow through
ecosystems below) tends to be less efficient during these early stages. The
number of trophic levels also tends to be lower. Early colonizers may change
the environment, facilitating later communities of other species to develop and
eventually become dominant species. The community of organisms at the final
stage of succession is called the climax community. These communities are
generally characterized by a wider range of species, a higher number of trophic
levels, and more efficient transfer of energy between trophic levels. In some
cases, a sequential process that changes the community of plants and animals
occurs in which there is initial colonization by fast growing, opportunistic
species followed by species that may grow more slowly but which compete

better for limited resources (light, moisture, nutrients). The final climax stage
reached is dependent on the pool of species available to colonize the area, and
the type, frequency, and intensity of initial disturbance (Christensen 1988).
Community succession, therefore, is dependent on both predictable and
stochastic events.
A successional series begins when some type of disturbance sets back the
clock. Landslides, hurricanes, avalanches, catastrophic forest fires, volcanic
eruptions, and meteorite impacts may set the clock back to zero by
completely destroying the existing communities. These types of events are
relatively rare and there are usually some species (e.g. fire-resistant seeds)
adapted to surviving these severe types of disturbances. Less severe
disturbances from localized storms, hurricanes, or burned forests are more
common and generally result in only partial damage to the community.
In coastal forests of South Carolina, including the ACE Basin, this progression
commonly starts with communities dominated by grasses and other fast
growing opportunistic weedy species that colonize the area within the first
few weeks or months after a disturbance. Over the next year or so, the
community is slowly taken over by woody shrubs and young pine trees that
grow more slowly than the opportunistic species. Over the next 20-30 years, as
the pines begin to grow large enough to shade the ground beneath, shadetolerant species may become established. Shade-intolerant plants that colonized
the open habitat, including young pine trees, can no longer establish themselves
under a closed canopy. Shade-tolerant species such as hardwood species (e.g.
beech and oak) become established, and the community develops into a
Southern mixed hardwood community. Between 50 and 100 years after the
disturbance, pine trees begin to slowly die, and the community gradually shifts
towards a hardwood dominated community, the common climax community of
coastal South Carolina upland areas. (See related section: Plants.)
Although fires and storms still may cause significant disturbance in ACE Basin
communities, humans impacts through agriculture and forestry practices and
urban development have far greater impact on succession in the ACE Basin.
One of these anthropogenic influences is the regulation of fire which may result
in changes in communities that are dependent on seasonal fires. Management
plans usually prescribe burns during the winter without regard to the natural
requirements of the species, because this is a period when areas are wetter and
fires are less likely to get out of control. This, in part, explains the reduced
dominance of longleaf pine forests that require spring and summer fires in the
early part of the growing season. Another impact is the clearing of fields (the
initial disturbance) and their maintenance as agricultural land (repeated

disturbances) preventing the successional sequence that would return them to a

forested landscape if left undisturbed. While some areas have been allowed to
develop back into forests, more frequently land has been shifted from forest to
agriculture and then on to suburban or urban uses. There are extreme cases,
fortunately outside the ACE Basin, where habitats have been so severely
impacted through salinization, erosion, or pollution that the areas would take
thousands of years to recover and may never revert to the habitat that was there
originally (Odum 1997). There is a field of study called restoration ecology that
deals with restoration and reclamation of severely degraded habitats.
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Biodiversity
One of the more important characteristics of a habitat is its biodiversity.
Individual species in a habitat fill different roles or niches including primary
production, herbivory, predation, and decomposition. In most communities
these roles are filled by many species with varied characteristics.
Biologically diverse communities, often occurring in stable environments
where complex assemblages of species have time to develop, can provide a
wide variety of services to the ecosystem as well as to humans interested in the
services. As an example, the potential medicinal value of the plants in the
biologically rich tropical regions is well recognized, and the loss of these
communities through deforestation may reduce this source of alternative
medicines. As another example, the loss of habitat for the Red-cockaded
woodpecker is directly related to current forest management practices of
harvesting pine forests before they have time to age to the point that
woodpeckers can utilize them.
The diversity of a community or habitat is, in part, dependent on the stability of
the environment surrounding the habitat. If environmental conditions remain
relatively constant, the available resources tend to be divided up between many
different species. The transfer of energy between trophic levels tends to be
more efficient in communities that have a number of trophic levels. These
conditions are characteristic of climax communities at the late stages
of succession. Examples of climax communities are the maritime forest
on barrier islandsand hardwood forests in inland areas that do not burn
regularly.

In environments where disturbances are relatively frequent, the diversity of

plant and animal species may be less. Frequent disturbance may keep the
development of the community in the early stages of succession, which tends to
have fewer species and a less efficient transfer of energy between trophic
levels. An example of this type of community is the pine flatwood community
where forest fires occur every 2-5 years, setting back the successional clock.
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Biological Populations
o Reproductive Rate
o Growth Rate and "r" and "k" Selected Species
o Carrying Capacity: How Many is Too Many?
The thousands of species and billions of individual plants and animals that
occupy the ACE Basin can be organized into groups by species
(populations), trophic levels (position in food chain), or habitats (communities).
The simplest level of organization of individuals is the population, a group of
individuals of one species living in a specific area. Every population has
attributes associated with it such as number of individuals (or density), range,
habitat requirements, reproductive rates, and dispersal characteristics.
Reproductive Rate
The reproductive rate of a population is the inherent or intrinsic growth rate of
a population in an unlimited environment. Most environments impose limits to
population growth, whether it be nutrients, space or water supply. However,
some habitats such as newly formed ponds or newly plowed fields may provide
enough nutrients and space that they are functionally unlimited for a short
period of time. In these cases, populations with high growth rates and a high
reproductive potential can experience explosive population growth and rapidly
colonize the habitat.
Growth Rate and r and k selected species
The growth rate of a population is a combination of the number of individuals
in the population, the death rate, the reproductive rate, and whatever limiting
factors act on the population. Growth rate or reproduction rate strategies can be

classified along a continuum between two endpoints. Species that are rselected have a high reproductive potential, producing many offspring. These
species, however, devote little energy to protecting their offspring and they
experience high mortality rates. Examples of organisms that are r-selected
include many plants that are considered weeds as well as many aquatic
worms and crustaceans that can rapidly colonize unoccupied habitats. In
general these species release large numbers of gametes with minimal parental
care. Species that are k-selected have a lower reproductive potential,
producing fewer offspring but devote much of their energy towards protecting
their offspring until they are ready to fend for themselves. These species may
have only one or two offspring every year, mature slowly, and are relatively
long-lived. The Florida manatee and humans are examples of k- selected
species, with low reproductive potential and long periods of parental care for
individual offspring. Other species fall somewhere along the continuum
between these two reproductive strategies.
In general, r-selected species tend to be opportunists, rapidly expanding their
populations in temporarily unlimited habitats. These species can rapidly
establish themselves in newly formed or recently disturbed habitats. After a
catastrophic forest fire for example, it is r-selected species that first colonize
the site. Similarly, the weeds that attempt to invade a corn field (disturbed
every year by site preparation) are usually r-selected species. The conditions
these species exploit are often characterized by an abundant resources with a
limited number of consumers. While able to rapidly invade an area, r-selected
species generally do not compete as well as k-selected species when
resources become limited. In the absence of frequent disturbances, r-selected
species are out-competed for limiting resources by k-selected species that
utilize limited resources more effectively. K- selected species tend to
dominate stable habitats such as climax communities, where competition for
space, light, and nutrients may be important.
Carrying capacity: How many is too many?
The carrying capacity of an environment is a theoretical maximum density of
individuals that a particular environment can support. In terrestrial habitats,
limiting factors are usually temperature, moisture regime, food, space, or
predation. In aquatic systems, especially in small streams and tidal creeks,
dissolved oxygen may also limit the number of individuals. Populations tend to
fluctuate around the carrying capacity, by exceeding the maximum carrying
capacity for short periods and then dropping below it. For food-limited
populations, as their population exceeds their food supply (exceeds the carrying
capacity) some individuals will starve or emigrate, thereby reducing the

population. The population falls and remains below the carrying capacity until
food is no longer limited. With adequate food resources, the population will
begin to increase again, repeating the pattern. In some cases, the fluctuations
are relatively small while other species may experience dramatic oscillations in
their population.
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What is a Community?
Any particular location supports populations of different species. Therefore, a
community is defined as all the different populations that live together in an
area. (Odum 1997). The area comprising the ACE Basin encompasses many
different communities. Communities can have unique boundaries, have varied
composition of species, and be variable in time and space. A community may
be defined as all the plants and animals (including humans) living on a
particular barrier island. Alternatively, a community may also be described in a
more specific sense, such as all the parasites living in or on a single fish such as
a spotted seatrout.
Different communities frequently have similar characteristics. They are usually
composed of both common and rare species. Some species, such as deer, are
generalists that live in a wide variety of habitats or feed on a wide range of
foods; others species are specialists such as the Red-cockaded woodpecker,
which requires specific microhabitats or food sources. Some of these specialists
are threatened by reductions of their required habitat. The Red-cockaded
woodpecker, an endangered species once found in the ACE Basin, requires pine
stands with a minimum age of 50-70 years. The loss of this habitat through
timber harvesting has resulted in the extirpation of Red-cockaded woodpeckers
from the ACE Basin. In contrast, the marsh hen is also a specialist, inhabiting
only the Spartina marshes. But, due to the expanse of this habitat in the ACE
basin, this species is very abundant there. National and state level restrictions
on impacting Spartina wetlands protect this habitat and therefore protect the
marsh hen.
External forces such as geomorphology (primarily affecting plant communities)
and episodic events such as storms or fire may also shape the structure of
communities. Low-lying areas are generally wet and support only those plant
species that can tolerate wet conditions. Sandy soils with limited organic

materials do not hold moisture well, and those habitats are dominated by
communities of plants and animals that can withstand dry conditions. Intense
storms such as hurricanes may decimate an Atlantic maritime forest dominated
by hardwoods allowing faster growing, shade intolerant, pine trees to establish
themselves and temporarily dominate the plant community.
In addition to external physical forces, a number of biological processes within
a community (such as species interactions) help to establish and maintain its
structure. Examples include territorial behavior, predator-prey interactions,
inhibitory chemicals used by plants to slow the growth of other plant species,
overstory shading that reduces the growth of shade intolerant species, and
parasites and diseases. All these processes interact to control the abundance and
diversity of the community of plants and animals in an area. Consider the
example of Edisto Beach: humans have greatly altered geomorphic features and
wetlands of the island. The structure of the community has changed
considerably through the modifications that humans have made to make the
island habitable for them. This may favor raccoons, egrets, and house cats at
the expense of salamanders and bald eagles.
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Energy and Trophic Levels

o Energy subsidies in Managed Systems
The energy supplied by the sun is the only source of truly renewable energy on
the earth. Other sources of energy, such as fossil fuels, are not renewable.
Nuclear energy, although potentially unlimited, has disadvantages related to its
waste products. Hydroelectric energy is renewable, but is dependent on solar
energy and weather patterns to replace the water that flows through the dam.
Most of the sunlight falling on the earth is absorbed as thermal energy (Energy
Dissipation of Solar Radiation as Percentage of Annual Input into the
Biosphere ) and this drives physical processes such as the hydrological cycle
through evaporation and rainfall, and weather patterns through the heating of
the air and land. Only about 1-5% of energy from the sun is stored
by photosynthesis while the rest is radiated as thermal energy or heat (Odum
1997).

Energy flow through the biosphere (the thin biological layer at the earths
surface) begins with the conversion of the radiant energy of sunlight to
chemical energy by organisms calledautotrophs. These include all the species,
such as plants, algae and some bacteria, that can convert solar energy to highenergy molecules through the process of photosynthesis. These species convert
raw materials such as elemental nitrogen, carbon dioxide, water, and potassium
into organic molecules, and in the process store energy in those molecules. The
organic materials synthesized in this way are used by autotrophs to grow and
survive. This conversion of raw materials and solar energy to organic materials
is called primary production. Gross primary production is the total amount of
raw materials converted by plants to organic molecules. A percentage of the
gross primary production is used to maintain the plants themselves. Any extra
production is called net primary production and it is used for growth and
reproduction or it may be lost as dissolved organic materials.
Energy stored by primary producers is used by primary consumers which
include herbivores and detritivores. Some herbivores include deer, rabbits,
seed-eating birds, caterpillars, and carp which feed on living plant material.
Detritivores are less conspicuous but no less important because they eat dead
plant material and start the decomposition processes that reduce the organic
material back to simpler, lower energy forms. From an energy storage and
transfer point of view, these levels of primary producer, primary consumer, and
secondary consumer are called trophic levels. Energy stored in high-energy
organic molecules is transferred between these levels by herbivores grazing on
plants and carnivores preying on herbivores. If it is hierarchical, from plant to
herbivore to carnivore, this flow of materials is called a food chain. However,
movement of materials from primary producers to consumers to decomposers
typically follows many different paths resulting in a complex network called
the food web as depicted in this figure from Odum (1997).
Energy Subsidies in Managed Systems
In a natural system that has been undisturbed by anthropogenic impacts, there
is a constant flow of energy, in the form of organic materials, that starts
with autotrophs using raw materials combined with solar energy to produce
high-energy organic molecules, which are then transferred up the food chain.
As plants and animals excrete wastes or die, decomposers break down these
waste materials, extracting the remaining energy from the organic molecules
and reducing the material back to raw elements. Almost all of the energy
needed to create this flow of energy is provided by solar energy with small
amounts coming from geothermal and chemical sources.

Humans have substantially changed the flow of energy in some systems by

subsidizing the energy input (Odum 1997). In the course of history, two major
changes in energy flow have allowed the human population to expand
significantly. The first was during the agricultural revolution, which occurred
approximately 8,000 years ago. The primary mechanisms of subsidizing energy
flow through agriculture became the addition of fertilizers (energy rich organic
material ) and removal of competitors (energy removed by unwanted sources
such as weeds and pests). The second change occurred with the industrial
revolution, when advances in medicines increased human life expectancies and
the use of fossil fuels increased the productivity of agriculture through
mechanization.
The global economy is currently using more energy than is renewable over the
long term and we are therefore heavily dependent on non-renewable fossil fuels
and nuclear energy. Our technologically advanced societies are using more
energy than can be replaced by renewable energy sources. Thus, energy use is
currently subsidized by fossil fuels, a limited resource; and nuclear energy an
unlimited resource that has a significant impact on the environment through the
release and disposal of radioactive byproducts. Renewable energy sources such
as hydroelectric power and solar power, although cleaner, have limitations as
well as undesirable impacts on the environment.
Energy use at the local level of the ACE Basin is no exception. Millions of
dollars in fuel, fertilizer, and pesticides are used to increase the production of
agricultural products well beyond natural levels. When fossil fuels become
expensive and scarce, how will the current production levels be maintained on
ACE Basin farms? One of the problems faced by society is how to maintain
current production levels as non-renewable resources such as fossil fuels
become scarce. This issue is not restricted just to the agricultural community
but will impact many of the ways that human society uses fossil fuels to
subsidize ecosystem services.
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Ecosystem Services
A Global Perspective

Ecosystem services are defined by Daily (1997) as the conditions and

processes through which natural ecosystems, and the species that make them
up, sustain and fulfill human life. Humans benefit from the materials and
processes provided by these services. Some of these materials such as seafood
and lumber are clearly apparent while others are less conspicuous including
waste assimilation, carbon cycling, storm buffering, and the hydrological cycle.
From a global perspective, ecosystems within the ACE Basin contribute to the
maintenance of the atmosphere by removing carbon dioxide and producing
oxygen, by transforming nitrogenous wastes into less toxic forms, and by
trapping pollutants. Global ecosystem services (Ecosystem services and
functions ) are large-scale processes that remain stable over long periods of
time, thereby allowing biological systems to develop. Loss of these ecosystem
services would result in significant changes in the stability of the global
ecosystem, affecting all life on earth. However, the scale and complexity of the
global ecosystem make it exceedingly difficult to detect changes; to attribute
causes for the change; and to stop, or reverse, the processes leading to the
change.
ACE Basin Perspective
The ACE Basin ecosystem is an integrated network of habitats that exchange
nutrients, decompose organic detritus, convert chemicals from organic
and inorganic states, capture energy, provide food and shelter, detoxify
pollutants, and provide economic opportunities. Some of these services, such as
food production, are readily apparent and have a market value. In the ACE
Basin, commercial fishing is an important means of food production. Likewise,
both agriculture and forestry products are produced in the ACE Basin and have
a market value. Less apparent services include biological and chemical
processes, such as the transfer of energy through the food chain, that operate to
produce the fish or agricultural products (Peterson and Lubchenco 1997; Odum
1997). These services are generally taken for granted, yet they may be severely
impacted by land use and pollution.
As an example, estuarine ecosystems absorb certain anthropogenic wastes
(such as nitrogen released from municipal waste-treatment plants and fertilizers
applied to agricultural fields) as well as trap and detoxify pollutants. However,
if these services are overloaded or the capacity of the estuary to process these
nutrients is reduced through wetland loss, the free service provided by the
estuary will have to be replaced by expensive technological substitute services
such as more efficient and expensive waste treatment plants.

Another example involves the dunes and maritime forests of the ACE Basin
which provide protection from storms. Dune lines serve to buffer coastal areas
from high seas and water levels during episodic events such as strong storms
and hurricanes. Human activities such as building on dunes, changing the
movement of sand with groins or jetties, or removing large areas of maritime
forest may compromise this protective buffer allowing increased erosion or
greater property damage (Thieler and Bush 1991).
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Wetlands as Contaminant Filters

Wetlands and estuaries trap many pollutants such as oil (PAHs), pesticides
(DDT, Chlordane), and heavy metals (lead, copper) in their sediments. Some of
these pollutants, such as PAHs, may then be further degraded by natural
microbial activity to less toxic byproducts or to simple compounds such as
carbon dioxide and water (Peterson and Lubchenco 1997). Pollutants that are
not readily degraded (PAHs, chlorinated compounds, and metals) are often
bound up with the fine particulates that make up the sediments and therefore
are less likely to be absorbed by wildlife. Sediments may temporarily trap the
pollutants, but biogenic and oxidative processes periodically facilitate the
release of sediment-associated pollutants. Therefore, pollutants may cause
long-term effects on biological resources of the ACE Basin. Although not yet
identified as a problem in the ACE Basin, many highly urban areas on the east
coast of the United States have exceeded the capability of these services, and
are experiencing the undesirable effects of these contaminants being released
into the environment and affecting wildlife as well as human populations.
These effects are seen in changes in plant and animal populations, increases in
fish mortality, reduced growth and deformities, and harmful algal blooms
(SCDHEC 1998). (See related section: Phytoplankton: Algal Blooms.)
Some ecosystem services can be difficult to assign a monetary value.
Ecologists and economists have made some recent progress in assigning dollar
values to ecosystem (Value of biome types ) operations or functions to
elucidate the economic value of pertinent non-market goods (Colgon 1990).
While these estimates are considered rough ones and are thought to
underestimate any actual dollar value equivalent of aggregate services of the
biome (Costanza et al. 1997), they make the value of these services comparable
with more easily recognized services such as the value of commercial fisheries

(Total value of commercial fishing in South Carolina.

aquatic systems to recycle municipal waste.

) or the ability of

Organisms are divided into autotrophs and heterotrophs according

to their energy pathways. Autotrophs are those organisms that are
able to make energy-containing organic molecules from inorganic
raw material by using basic energy sources such as sunlight. Plants
are the prime example of autotrophs, using photosynthesis. All other
organisms must make use of food that comes from other organisms
in the form of fats, carbohydrates and proteins. These organisms
which feed on others are called heterotrophs.

A heterotroph (/htrtrof/; heteros = "another", "different"

and trophe = "nutrition") is an organismthat cannot fix carbon and
uses organic carbon for growth.[1][2] Heterotrophs can be further divided based
on how they obtain energy; if the heterotroph uses light for energy, then it is
considered a photoheterotroph, while if the heterotroph uses chemical energy,
it is considered a chemoheterotroph.
Heterotrophs contrast with autotrophs, such as plants and algae, which can
use energy from sunlight(photoautotrophs) or inorganic compounds
(lithoautotrophs) to produce organic compounds such ascarbohydrates, fats,
and proteins from inorganic carbon dioxide. These reduced carbon
compounds can be used as an energy source by the autotroph and provide
the energy in food consumed by heterotrophs. Ninety-five percent or more of
all types of living organisms are heterotrophic.[3]
An autotroph[] ("self-feeding", from the Greek autos "self"
and trophe "nourishing") or "producer", is an organism that produces
complex organic compounds (such as carbohydrates, fats, and proteins) from
simple substances present in its surroundings, generally using energy from
light (photosynthesis) or inorganic chemical reactions (chemosynthesis). They
are the producers in a food chain, such as plants on land or algae in water, in
contrast toheterotrophs as consumers of autotrophs. They do not need a living
energy or organic carbon source. Autotrophs can reduce carbon dioxide to
make organic compounds for biosynthesis and also create a store of chemical
energy. Most autotrophs use water as the reducing agent, but some can use
other hydrogen compounds such as hydrogen sulfide. Phototrophs (green
plants and algae), a type of autotroph, convert electromagnetic energy from
sunlight into chemical energy in the form of reduced carbon.
Autotrophs can be photoautotrophs or chemoautotrophs. Phototrophs use
light as an energy source, while chemotrophs utilize electron donors as a
source of energy, whether from organic or inorganic sources; however in the
case of autotrophs, these electron donors come from inorganic chemical

sources. Such chemotrophs arelithotrophs. Lithotrophs use inorganic

compounds, such as hydrogen sulfide,
elemental sulfur, ammonium andferrous iron, as reducing agents
for biosynthesis and chemical energy storage. Photoautotrophs and
lithoautotrophs use a portion of the ATP produced during photosynthesis or
the oxidation of inorganic compounds to reduce NADP+ to NADPH to form
organic compounds.[1]
Herbivores

Herbivores are animals which only eat plant material. This means leaves, flowers, fruits
or even wood. Sheep, horses, rabbits and snails are well known examples of herbivores
which eat grass and leaves. A parrot, however, which eats fruits and nuts can also be
called a herbivore.
Omnivores

Omnivores eat both plants and meat. Chickens are omnivores. They eat seeds, but they
can also eat worms. human beings are also omnivores, although some people choose
not to eat meat. These people are called vegetarians.
The chimpanzee is omnivorous. It eats fruits, leaves, palm nuts, seeds and stems, as
well as ants, birds' eggs, fish and termites. Chimpanzees will occasionally kill and eat
baboons and wild pigs.
Carnivores

Carnivores eat meat. A carnivore is a predator because it has to find and catch its prey.
Some carnivores, such as wolves, hunt in a group called a pack. They move silently and
slowly to form a circle around their prey before they attack.
Johnny's Fact File No.6
The smallest carnivorous mammal is called the least weasel. This little mammal is
found in North America. It's mass is 57 grammes at the most!

Other carnivores, such as the cheetah, usually hunt alone. The cheetah creeps towards
its prey without being noticed, until it is 30 metres from it, then the cheetah starts to run.
Some insects are carnivores. The dragonfly, which hovers so gracefully above a pond,
is hunting for other insects.
The eagle is a carnivorous bird. It flies high in the sky looking for animals, such as
rabbits. When it finds one it quickly swoops to the ground. It uses its strong feet and
pointed claws, called talons, to catch the rabbit. It has a very pointed beak to help it tear
off the meat.