MODULE 5

Community ecology

Learning Outcomes:
After studying this chapter, students are expected to:
1. Examine the process of synecology or community ecology.
2. Understand theoretical models suggesting relationship between diversity and
ecosystem stability.
3. Recall pioneering studies of scientific experiment on diversity-stability
relationship.
4. Understand the concept of island biogeography or insular biogeography.
5. Integrate processes of community change and assemblage.
6. Classify the kinds of ecological communities or biomes of terrestrial and
aquatic ecosystem.

On a deeper level the meaning and value of the community concept in ecology is up for
debate. Communities have traditionally been understood on a fine scale in terms of local
processes constructing (or destructing) an assemblage of species, such as the way climate change
is likely to affect the make-up of grass communities. Recently this local community focus has
been criticized. Robert Ricklefs has argued that it is more useful to think of communities on a
regional scale, drawing on evolutionary taxonomy and biogeography, where some species or
clades evolve, and others go extinct.
Community ecology or synecology is the study of the interactions between species in
communities on many spatial and temporal scales, including the distribution, structure,
abundance, demography, and interactions between coexisting populations. The primary focus of
community ecology is on the interactions between populations as determined by specific
genotypic and phenotypic characteristics.
Community ecology also takes into account abiotic factors e.g. annual temperature or soil
pH. These non-living factors can influence the way species interact with each other. Abiotic
factors filter the species that are present in the community, and therefore community structure.
For example, the difference in plants present in the desert compared to the tropical rainforest is
dictated by the annual precipitation. These non-living factors also influence the way species
interact with each other. Humans can also have an effect on community structure through habitat
disturbance, such as introduction of invasive species.
Community ecology has its origin in European plant sociology. It examines processes
such as predator–prey population dynamics or succession. Whilst also examining patterns such
as variation in:
 Species richness- is the number or simply a count of different species represented in an
 ecological community, landscape, or region.
 Species evenness- refers to how close in numbers each species in an environment is.
 Biodiversity- is the variety and variability of life on earth.
 Productivity (ecology)- refers to the rate of generation or biomass in an ecosystem,
usually expressed in units of mass per volume (unit surface) per unit of time, such as
grams per square meter per day (g m-2d-1)
 Food web- is the natural interconnection of food chain and a graphical representation
(usually an image) of what-eats-what in an ecological community.
 Community structure- the study of complex networks, a network is said to have
community structure if the nodes of the network can be easily grouped into (potentially
overlapping) sets of nodes such that each set of node densely connected internally.

5.1 Species Diversity and Community Stability

Traditional ecological wisdom says Diversity Begats Stability (Charles Elton), meaning
more complex ecosystems have more checks and balances.

Diversity-Stability Theory
Theoretical models suggest that there could be
multiple relationships between diversity and stability,
depending on how we define stability . Stability can be
defined at the ecosystem level — for example, a rancher
might be interested in the ability of a grassland ecosystem to
maintain primary production for cattle forage across several
years that may vary in their average temperature and
precipitation. Figure 1 shows how having multiple species
present in a plant community can stabilize ecosystem
processes if species vary in their responses to environmental
fluctuations such that an increased abundance of one species
can compensate for the decreased abundance of another.
Biologically diverse communities are also more likely to
contain species that confer resilience to that ecosystem because
as a community accumulates species, there is a higher chance
of any one of them having traits that enable them to adapt to a
changing environment. Such species could buffer the system
against the loss of other species. Scientists have proposed the
insurance hypothesis to explain this phenomenon (Yachi &
Loreau 1999). In this situation, species identity — and particular species traits — are the driving
force stabilizing the system rather than species richness per se (see Figure 2).

Experiments and Observations Can Evaluate the Diversity-Stability Relationship

A wealth of research into the relationships among diversity, stability, and ecosystem
functioning has been conducted in recent years (reviewed by Balvanera et al. 2006, Hooper et al.
2005). The first experiments to measure the relationship between diversity and stability
manipulated diversity in aquatic microcosms — miniature experimental ecosystems —
containing four or more trophic levels, including primary producers, primary and secondary
consumers, and decomposers (McGrady-Steed et al. 1997, Naeem & Li 1997). These
experiments found that species diversity conferred spatial and temporal stability on several
ecosystem functions. Stability was conferred by species richness, both within and among
functional groups (Wardle et al. 2000). When there is more than one species with a similar
ecological role in a system, they are sometimes considered "functionally redundant." But these
experiments show that having functionally redundant species may play an important role in
ensuring ecosystem stability when individual species are lost due to environmental changes, such
as climate change.
More recently, scientists have examined the importance of plant diversity for ecosystem
stability in terrestrial ecosystems, especially grasslands where the dominant vegetation lies low
to the ground and is easy to manipulate experimentally. In 1995, David Tilman and colleagues
established 168 experimental plots in the Cedar Creek Ecosystem Science Reserve, each 9 x 9 m
in size (Figure 3A), and seeded them with 1, 2, 4, 8 or 16 species drawn randomly from a pool of
18 possible perennial plant species (Tilman et al. 2006). Plots were weeded to prevent new
species invasion and ecosystem stability was
measured as the stability of primary production over
time. Over the ten years that data were collected,
there was significant interannual variation in
climate, and the researchers found that more diverse
plots had more stable production over time (Figure
3B). In contrast, population stability declined in
more diverse plots (Figure 3C). These experimental
findings are consistent with the theory described in
the prior section, predicting that increasing species
diversity would be positively correlated with
increasing stability at the ecosystem-level and
negatively correlated with species-level stability due
to declining population sizes of individual species.
5.2 Island Biogeography
Theory of Island Biogeography
Island biogeography (also called insular biogeography)
provides some of the best evidence in support of natural
selection and the theory of evolution. The term describes an
ecosystem that is isolated by being surrounded by different
ecosystems. For the purposes of this theory, an island is defined
as more than just a piece of land surrounded by water. It
includes mountain peaks, a lake surrounded by a desert, a patch
of woodland or even a national park. The theory provides a
model to explain the richness and uniqueness of species, both
plants and animals, found in an isolated area.
The two events that determine how many species are
found in an isolated ecosystem are immigration and extinction.
Research as shown that how big the island is and how far it is
from the mainland have a great influence on the number (richness) of species that are found there
(see image below). Once species have established themselves on an island, the rate at which they
will go extinct depends on the size of the island, with there being less likelihood of extinction on
larger islands. This is called the species-area relationship. This relationship is not just observable,
but it can also be predicted mathematically. By the same theory, the farther an island is from the
mainland, the fewer species it tends to have. This is referred to as the species-distance
relationship.

Alfred Russel Wallace, the Father of Biogeography

In the mid-19th century, the British naturalist and explorer Alfred Russel Wallace traveled
to the Amazon Basin and the Malay Archipelago (located between China and Australia) to build
on the work of Charles Darwin and study how and why the flora and fauna were distributed. He
paid particular attention to the distributions of butterflies and birds in relation to the presence and
absence of natural geographic barriers like rivers, valleys and mountains. Wallace’s findings
contributed significant knowledge to the field of biogeography, including his discovery of the
sharp contrasting differences in the animals on either side of what is now the Wallace Line,
named to acknowledge the importance of his work.

Island Biogeography Examples:

Australia
Marsupials like the kangaroo and the wallaby are only found in Australia. If marsupials
were found all over the world, then that would mean they did not come into existence by means
of natural selection and the evolutionary process. Additional evidence for this evolutionary
model is shown by the fact that about 75% of the plant and mammal species in Australia are
found only on that continent. There are some exceptions, however, due to continental drift when
animals were able to wander around large land masses before they separated.

The Galapagos and Cape Verde Islands

When he visited the Galapagos and Cape Verde Islands in 1835, Charles Darwin
encountered species that are found nowhere else on Earth like the Galapagos Tortoise, the
Flightless Cormorant, the Blue-footed Boobie and, of course, the famous Darwin’s Finches. But
it’s not just the fact that these isolated environments gave rise to unique species—each island
also had its own unique species. In addition, Darwin noted at the time that none of the species
from these islands were found in similar climates anywhere in the world. Darwin wondered why
there were distinct species on each of the islands when the climates are virtually identical, and
why there appeared to be closely related species on the nearest mainland continent. These
observations led to his hypothesis (at the time) that the islands had broken off separately from the
main continent sometime in the distant past, resulting in two identical populations that evolved
separately over time

5.3 Community Change

TOPICS
 Ecology and Time’s Environmental Texture
 Communities Come, Communities Go
 What Governs Community Assembly and Disassembly?
 Spanning the Missing Middle
 Community Assembly and Disassembly.
 Trait-Based Community Patterns.
 Diversity Dynamics Through Time.
 Ecological Rules in the Anthropocene.
 Dynamics of Regional Species Pools.
 Clocking Time-Dependent Processes.
  Meeting Cowles’ Challenge

Community ecology and paleoecology are both concerned with the composition and
structure of biotic assemblages but are largely disconnected. Community ecology focuses on
existing species assemblages and recently has begun to integrate history (phylogeny and
continental or intercontinental dispersal) to constrain community processes. This division has left
a “missing middle”: Ecological and environmental processes occurring on timescales from
decades to millennia are not yet fully incorporated into community ecology. Quaternary
paleoecology has a wealth of data documenting ecological dynamics at these timescales, and
both fields can benefit from greater interaction and articulation. We discuss ecological insights
revealed by Quaternary terrestrial records, suggest foundations for bridging between the
disciplines, and identify topics where the disciplines can engage to mutual benefit.

Ecology and Time’s Environmental Texture

Much of ecological concept and theory has considered time as a simple, uniform
dimension, wherein ecological processes unfold against a constant environmental backdrop.
Time is treated as an independent variable, and ecological properties change as time-dependent
ecological processes run their course. A classic example is the exponential model of population
growth, together with its many derivative models incorporating resource competition, predation,
multiple species, age structure, time lags, and other features. The environment itself may change,
but only as a consequence of the ecological processes themselves (e.g., consumption of
resources; autogenic successional processes such as soil development or lake infilling) or as a
stochastic variable, fluctuating randomly about a constant mean. Long-term environmental
change has long been acknowledged by ecologists, but only recently have environmental change
and nonstationary variability been explicitly incorporated into ecological models, usually as
discrete perturbations or at relatively short time intervals (seasons to decades).

Communities Come, Communities Go

Ecological community comprises a single place that happens to be occupied by an
assemblage of species with overlapping distributions and environmental tolerances. It is argued
that the community concept be “replaced by the spatial distributions of populations, which now
become the primary focus for understanding biodiversity patterns”. Paleoecological records
support a parallel conception involving time. An ecological community can be viewed as a single
point in a spatial framework of species distributions superimposed on environmental gradients
and patchworks and in a temporal framework of population and biogeographic responses to
environmental change and variability. The space and time dimensions cannot be treated
separately; spatial distributions evolve through time and are often contingent on temporal
processes. Conversely, the temporal patterns of occurrence and abundance at an individual site
are contingent on spatial distributions, patterns, and processes. Understanding of community
properties is inevitably incomplete in the absence of both spatial and temporal context.

What Governs Community Assembly and Disassembly?

Determinants of community structure and composition are a central focus of community
ecology and tend to align with one of three modal concepts: interaction assembly, environment
assembly, or neutral assembly . Interaction assembly is deeply rooted in community ecology and
emphasizes the Eltonian niche, considering communities to be structured primarily by strong
interactions among species. Such interactions may include facilitation, mutualism, and trophic
relationships (predation, parasitism, herbivory), but resource competition often is treated as first
among equals. Interaction-based communities have limited membership, governed by
equilibrium population processes as determined by interspecific interactions. In contrast, neutral
assembly considers communities to be structured entirely by random processes, particularly
dispersal, recruitment, and mortality. Neutral communities have virtually unlimited membership,
are nonequilibrium, and bear historical imprints; composition is influenced by legacies of past
demographic and dispersal events, and hence community properties may drift as the result of
singular events. Finally, communities under environment assembly are structured primarily by
species’ physiological and demographic responses to the physical environment. The
environment-assembly concept is also niche-based but emphasizes the Grinnellian niche. It is
predicated on all species having finite environmental requirements or tolerances, which impose
strong filters on community membership. Community composition is governed by whether
potential members’ fundamental (Grinnellian) niches overlap with the local environmental
realization. These three concepts are not mutually exclusive, and some efforts have been made to
reconcile interaction assembly and neutral assembly. Environment assembly has received less
attention in community ecology than the other two, although it underlies the recent explosion of
empirical models predicting species distribution and abundance and community composition
under various climate-change scenarios.

Spanning the Missing Middle

The missing middle of Quaternary ecological history provides abundant opportunity for
collaboration between paleoecologists and community ecologists. Success will require mutual
understanding and engagement. Paleoecologists must understand the ecological questions to
which their data are applied, and ecologists must understand the nature of paleoecological data
and inference. To avoid mismatches between data and applications, careful attention must be
paid to scale and taphonomy. We identify a nonexhaustive set of themes, topics, and questions
upon which collaborations might be centered.
Community Assembly and Disassembly.
Paleoecological records show not only that communities come and go but also that
community turnover occurs in many forms and at many rates. Some communities arise quickly,
whereas others develop in slow transitions. Conversely some communities disassemble gradually
via decline and replacement of dominants, but others appear to undergo rapid collapse. These
and other records provide diverse case studies for understanding the various roles and
interactions of environmental forcing, intrinsic community properties (e.g., resilience, hysteresis,
keystone species), and ecological drift in community transitions, whether gradual or abrupt.
Specific critical transitions can be targeted for intensive study, based on various criteria (e.g.,
knowledge of natural history of taxa, existence of independent paleoclimatic records, precision
of paleo ecological records, and availability of multiple paleo ecological sites for replication or
pattern analysis, potential significance of observed patterns). The rapid increase or decline of a
dominant species or the disappearance of a community, for example, is of obvious interest in
conservation context and may be driven by rapid environmental change, cross-scale interactions,
or both. Intensive, integrated study of a carefully selected array of paleoecological case studies
would test fundamental theory and indicate the extent to which community turnover follows
general rules. In addition, paleoecological data from individual sites or site arrays can be
analyzed to test specific theoretical predictions concerning community turnover.

Trait-Based Community Patterns.

Trait-based assembly and inference have become an important focus in community
ecology. Traits and related functional groups of species may govern whether and in what order
species can join communities, and influence interactions of individuals with the environment and
with each other. Paleontology has a rich tradition of connecting traits with ecological and
evolutionary processes, and trait-based approaches to community assembly are ripe for
paleoecological application. A critical challenge is to determine which traits extractable from the
fossil record map directly or indirectly onto niche properties (Grinnellian or Eltonian), and how
they are related to traits commonly measured by modern ecologists. An emerging agenda in
paleoecology is to track specific traits through time with the goal of explicitly connecting the
past with the present. Merging trait-based paleoecology with community ecology can identify the
stability of trait relationships through time. Much like temporal variation in species and
communities, trait distributions within species are likely to vary through time and with changing
environments. Paleoecological trait studies can assess conservatism of trait–environment
relationships and trait properties within taxa. Furthermore, they can help identify the extent to
which communities may display functional stasis while undergoing compositional turnover.

Diversity Dynamics Through Time.

 A growing body of work has demonstrated that environmental variability affects species
diversity on short time scales , suggesting a species–time relationship as an analog to the
species–area relationship. However, key differences between spatial and temporal processes
imply that they may have unique diversity-scaling relationships. In addition, space and time
constrain one another, motivating theoretical formulation and empirical testing of integrated
species–time–area relationships. White et al. (103) point to several fruitful areas to promote this
integration. However, most of their case studies are modern, encompassing climatic variation
over a few decades at most (e.g., refs. 101, 105), and their single fossil example starts with an
interval length of >700,000 y . The missing middle remains unfilled. Brown et al. (102)
examined how environmental variability may regulate diversity dynamics in several systems,
including Holocene fossil pollen records at family resolution. They postulated that although
environmental change maintains diversity by influencing colonization and extinction of different
species, when sufficiently large it could alter diversity by changing species-level carrying
capacity. These conjectures remain unexplored.

Ecological Rules in the Anthropocene.

Have ecological and evolutionary processes changed fundamentally during the
Anthropocene (107)? Do anthropogenic activities override past controls on community
dynamics? Addressing these questions requires accurate baselines to compare diversity trends
and patterns, and the Quaternary provides critical linkages between deep time and the
Anthropocene. For example, mammalian biodiversity patterns throughout the past 30 million
years suggest that the Holocene is different from all time periods that came before , implying
human impacts well before the Industrial Revolution. In some settings, beta diversity and
functional diversity appear to have remained relatively stable despite compositional changes,
providing context to evaluate whether ongoing processes override those of the past few million
of years. Beyond issues of the Anthropocene, there are legitimate questions concerning whether
the Quaternary itself is unique and whether ecological rules evolve with the global
environmental setting . Existing paleoecological records provide an untapped source for
investigation of diversity dynamics in diverse systems and at different timespans .

Dynamics of Regional Species Pools.

Community ecologists are focusing attention on the biogeographic and phylogenetic
contexts of species pools and their implications for community properties. These studies tend to
treat the regional species pool as static, subject to the same processes as (sub)continental pools.
Regional species pools, however, are subject to at least partial turnover over hundreds to
thousands of years and are predicted to change rapidly under global change because of both
extinction and range shifts. Implications of species-pool dynamics are being explored from
biogeographical and macroecological perspectives, but the consequences for community
assembly and structure represent open territory for collaboration between community ecologists
and paleoecologists.

Clocking Time-Dependent Processes.

Many ecological processes are time-dependent, and paleoecological studies indicate that
similar plant communities are established at different times in different places. Comparative
studies of secondary communities (mycorrhizae, invertebrates, vertebrates), using antiquity of
the plant community as an independent variable, can provide insights into processes of
community development and evolution. Age may be one among many differences among sites,
but other properties (e.g., physical environment, climate history, genetics) can be controlled in
study design. Time-dependent processes may be sufficiently large to override other effects, many
of which can be assessed directly from paleoecological and paleoenvironmental studies. For
example, ancient DNA may provide information on local genetic changes.

Meeting Cowles’ Challenge

More than a century ago Henry Chandler Cowles identified the central challenge of
ecological dynamics, observing that vegetational response to the environment was “a variable
approaching a variable rather than a constant”. The challenge has only grown since Cowles’
time: Ecological dynamics comprise a multitude of variables, with different response times,
changing at different rates, and often interacting in subtle ways. The environment, too, is far
more dynamic than envisioned even a decade ago, with nested climatic changes interacting
across a range of temporal and spatial scales. Community ecology offers tools and theory to help
understand ecological change; paleoecology offers diverse data and observations of actual
ecological change. By working together, these disciplines can advance understanding of
ecological dynamics in changing environments.

5.4 Ecosystems (Main Types of Communities)

Ecologists find that within a community many populations are not randomly distributed.
This recognition that there was a pattern and process of spatial distribution of species was a
major accomplishment of ecology. Two of the most important patterns are open community
structure and the relative rarity of species within a community.

Classification of Communities
 Image from: Purves et.al There are two basic categories of
communities: terrestrial (land) and aquatic
(water). These two basic types of community
contain eight smaller units known as biomes. A
biome is a large-scale category containing many
communities of a similar nature, whose
distribution is largely controlled by climate.

 Terrestrial Biomes: tundra, grassland, desert, taiga, temperate forest, tropical forest.
Terrestrial biome.

 Aquatic Biomes: marine, freshwater.

Terrestrial Biomes
 Tundra and Desert
The tundra and desert biomes occupy the most extreme environments, with little or no
moisture and extremes of temperature acting as harsh selective agents on organisms that
occupy these areas. These two biomes have the fewest numbers of species due to the
stringent environmental conditions. In other words, not everyone can live there due to the
specialized adaptations required by the environment.

 Tropical Rain Forests

Tropical rain forests occur in regions near the equator. The climate is
always warm (between 20° and 25° C) with plenty of rainfall (at least
190 cm/year). The rain forest is probably the richest biome, both in
diversity and in total biomass. The tropical rain forest has a complex
structure, with many levels of life. More than half of all terrestrial
Botanical society
species live in this biome. While diversity is high, dominance by a
of America particular species is low.

 Temperate Forests
The temperate forest biome occurs south of the taiga in eastern North
America, eastern Asia, and much of Europe. Rainfall is abundant (30-
80 inches/year; 75-150 cm) and there is a well-defined growing season
of between 140 and 300 days. The eastern United States and Canada
are covered (or rather were once covered) by this biome's natural

Image: Botany.org
vegetation, the eastern deciduous forest. Dominant plants include beech, maple, oak; and
other deciduous hardwood trees. Trees of a deciduous forest have broad leaves, which they
lose in the fall and grow again in the spring.
 Shrubland (Chaparral)
The shrubland biome is dominated by shrubs with small but thick
evergreen leaves that are often coated with a thick, waxy cuticle,
and with thick underground stems that survive the dry summers
and frequent fires. Shrublands occur in parts of South America,
western Australia, central Chile, and around the Mediterranean
Sea. Dense shrubland in California, where the summers are hot
and very dry, is known as chaparral, shown in Figure 4. This
Mediterranean-type shrub land lacks an understory and ground
Image: botany.org litter, and is also highly flammable. The seeds of many species
require the heat and scarring action of fire to induce germination.
 Grasslands
Grasslands occur in temperate and tropical areas with reduced rainfall (10-30 inches per
year) or prolonged dry seasons. Grasslands occur in the Americas, Africa, Asia, and
Australia. Soils in this region are deep and rich and are excellent for agriculture. Grasslands
are almost entirely devoid of trees, and can support large herds of grazing animals. Natural
grasslands once covered over 40 percent of the earth's land surface. In temperate areas where
rainfall is between 10 and 30 inches a year, grassland is the climax community because it is
too wet for desert and too dry for forests.

 Deserts
Deserts are characterized by dry conditions (usually less than 10 inches per year; 25 cm) and
a wide temperature range. The dry air leads to wide daily temperature fluctuations from
freezing at night to over 120 degrees during the day. Most deserts occur at latitudes of 30o N
or S where descending air masses are dry. Some deserts occur in the rainshadow of tall
mountain ranges or in coastal areas near cold offshore currents. Plants in this biome have
developed a series of adaptations (such as succulent stems, and small, spiny, or absent leaves)
to conserve water and deal with these temperature extremes. Photosynthetic modifications
(CAM) are another strategy to life in the drylands.
 Taiga (Boreal Forest)

The taiga (pronounced "tie-guh") is a coniferous forest extending

across most of the northern area of northern Eurasia and North
America. This forest belt also occurs in a few other areas, where it

Image: botany.org
 has different names: the montane coniferous forest when near mountain tops; and the
temperate rain forest along the Pacific Coast as far south as California. The taiga receives
between 10 and 40 inches of rain per year and has a short growing season. Winters are cold
and short, while summers tend to be cool. The taiga is noted for its great stands of spruce, fir,
hemlock, and pine. These trees have thick protective leaves and bark, as well as needlelike
(evergreen) leaves can withstand the weight of accumulated snow. Taiga forests have a
limited understory of plants, and a forest floor covered by low-lying mosses and lichens.
Conifers, alders, birch and willow are common plants; wolves, grizzly bears, moose, and
caribou are common animals. Dominance of a few species is pronounced, but diversity is low
when compared to temperate and tropical biomes.
 Tundra
The tundra, covers the northernmost regions of
North America and Eurasia, about 20% of the
Earth's land area. This biome receives about 20 cm
(8-10 inches) of rainfall months. Winters are long
and dark, followed by very short summers. Water is
frozen most of the time, producing frozen soil,
Alaska Conservation Foundation permafrost. Vegetation includes no trees, but rather
patches of grass and shrubs; grazing musk ox, reindeer,
and caribou exist along with wolves, lynx, and rodents. A few animals highly adapted to cold
live in the tundra year-round (lemming, ptarmigan). During the summer the tundra hosts
numerous insects and migratory animals. The ground is nearly completely covered with
sedges and short grasses during the short summer. There are also plenty of patches of lichens
and mosses. Dwarf woody shrubs flower and produce seeds quickly during the short growing
season. The alpine tundra occurs above the timberline on mountain ranges, and may contain
many of the same plants as the arctic tundra.

Aquatic Biomes

Like terrestrial biomes, aquatic biomes are

influenced by a series of abiotic factors. The aquatic
medium—water— has different physical and
chemical properties than air. Even if the water in a
pond or other body of water is perfectly clear (there
are no suspended particles), water still absorbs light.
As one descends into a deep body of water, there will
eventually be a depth which the sunlight cannot
 reach. While there are some abiotic and biotic factors in a terrestrial ecosystem that might
obscure light (like fog, dust, or insect swarms), usually these are not permanent features of
the environment. The importance of light in aquatic biomes is central to the communities of
organisms found in both freshwater and marine ecosystems. In freshwater systems,
stratification due to differences in density is perhaps the most critical abiotic factor and is
related to the energy aspects of light. The thermal properties of water (rates of heating and
cooling) are significant to the function of marine systems and have major impacts on global
climate and weather patterns. Marine systems are also influenced by large-scale physical
water movements, such as currents; these are less important in most freshwater lakes.
The ocean is categorized by several areas or zones (Figure 1). All of the ocean’s open
water is referred to as the pelagic zone. The benthic zone extends along the ocean bottom
from the shoreline to the deepest parts of the ocean floor. Within the pelagic realm is the
photic zone, which is the portion of the ocean that light can penetrate (approximately 200 m
or 650 ft). At depths greater than 200 m, light cannot penetrate; thus, this is referred to as the
aphotic zone. The
majority of the ocean is aphotic and lacks sufficient light for photosynthesis. The deepest part
of the ocean, the Challenger Deep (in the Mariana Trench, located in the western Pacific
Ocean), is about 11,000 m (about 6.8 mi) deep. To give some perspective on the depth of this
trench, the ocean is, on average, 4267 m. These zones are relevant to freshwater lakes as
well.

Marine Biomes
 Ocean
The ocean is the largest marine biome. It is a continuous body of salt water that is relatively
uniform in chemical composition; it is a weak solution of mineral salts and decayed
biological matter. Within the ocean, coral reefs are a second kind of marine biome. Estuaries,
coastal areas where salt water and fresh water mix, form a third unique marine biome. The
physical diversity of the ocean is a significant influence on plants, animals, and other
organisms. The ocean is categorized into different zones based on how far light reaches into
the water. Each zone has a distinct group of species adapted to the biotic and abiotic
conditions particular to that zone.
 Coral Reefs
Coral reefs are characterized by high biodiversity and the structures created by
invertebrates that live in warm, shallow waters within the photic zone of the ocean. They
are mostly found within 30 degrees north and south of the equator. The Great Barrier
Reef is a well-known
 reef system located several miles off the northeastern
coast of Australia. The coral organisms (members of
phylum Cnidaria) are colonies of saltwater polyps
that secrete a calcium carbonate skeleton. These
calcium-rich skeletons slowly accumulate, forming
the underwater reef (Figure 3). Corals found in
shallower waters (at a depth of approximately 60 m
or about 200 ft) have a mutualistic relationship with
photosynthetic unicellular algae. The relationship
provides corals with the majority of the nutrition and
the energy they require. The waters in which these
corals live are nutritionally poor and, without this
mutualism, it would not be possible for large corals
to grow. Some corals living in deeper and colder
water do not have a mutualistic relationship with
algae; these corals attain energy and nutrients using
stinging cells on their tentacles to capture prey. It is
estimated that more than 4,000 fish species inhabit coral reefs. These fishes can feed on
coral, other invertebrates, or the seaweed and algae that are associated with the coral.

 Estuaries: Where the Ocean Meets Fresh Water

Estuaries are biomes that occur where a source of fresh water, such as a river, meets the
ocean. Therefore, both fresh water and salt water are found in the same vicinity; mixing
results in a diluted (brackish) saltwater. Estuaries form protected areas where many of the
young offspring of crustaceans, mollusks, and fish begin their lives. Salinity is a very
important factor that influences the organisms and the adaptations of the organisms found in
estuaries. The salinity of estuaries varies and is based on the rate of flow of its freshwater
sources. Once or twice a day, high tides bring salt water into the estuary. Low tides occurring
at the same frequency reverse the current of salt water.

 Freshwater Biomes
Freshwater biomes include lakes and ponds (standing water) as well as rivers and streams
(flowing water). They also include wetlands, which will be discussed later. Humans rely on
freshwater biomes to provide aquatic resources for drinking water, crop irrigation, sanitation,
and industry. These various roles and human benefits are
referred to as ecosystem services. Lakes and ponds are found in
terrestrial landscapes and are, therefore, connected with abiotic
and biotic factors influencing these terrestrial biomes.

 Lakes and Ponds

Lakes and ponds can range in area from a few square meters to thousands of square
kilometers. Temperature is an important abiotic factor affecting living things found in lakes
and ponds. In the summer, thermal stratification of lakes and ponds occurs when the upper
layer of water is warmed by the sun and does not mix with deeper, cooler water. Light can
penetrate within the photic zone of the lake or pond. Phytoplankton (small photosynthetic
organisms such as algae and cyanobacteria that float in the water) are found here and carry
out photosynthesis, providing the base of the food web of lakes and ponds. Zooplankton
(very small animals that float in the water), such as rotifers and small crustaceans, consume
these phytoplankton. At the bottom of lakes and ponds, bacteria in the aphotic zone break
down dead organisms that sink to the bottom.

 Rivers and Streams

Rivers and streams are continuously moving bodies of water that carry large amounts of
water from the source, or headwater, to a lake or ocean. The largest rivers include the Nile
River in Africa, the Amazon River in South America, and the Mississippi River in North
America.
Abiotic features of rivers and streams vary along the length of the river or stream. Streams
begin at the point of origin referred to as source water. The source water is usually cold, low
in nutrients, and clear. The channel (the width of the river or stream) is narrower than at any
other place along the length of the river or stream. Because of this, the current is often faster
here than at any other point of the river or stream.
 Wetlands
Wetlands are environments in which the soil is either
permanently or periodically saturated with water. Wetlands
are different from lakes because wetlands are shallow
bodies of water that may periodically dry out. Emergent
vegetation consists of wetland plants that are rooted in the
soil but have portions of leaves, stems, and flowers
extending above the water’s surface. There are several
types of wetlands including marshes, swamps, bogs,
mudflats, and salt marshes