Biodiversity, a contraction of the phrase "biological diversity," is a complex topic,

covering many aspects of biological variation. In popular usage, the word biodiversity is
often used to describe all the species living in a particular area. If we consider this area at
its largest scale - the entire world - then biodiversity can be summarized as "life on
earth." However, scientists use a broader definition of biodiversity, designed to include
not only living organisms and their complex interactions, but also interactions with the
abiotic (non-living) aspects of their environment. Definitions emphasizing one aspect or
another of this biological variation can be found throughout the scientific and lay
literature (see Gaston, 1996: Table 1.1). For the purposes of this module, biodiversity is
defined as:

the variety of life on Earth at all its levels, from genes to ecosystems, and the ecological
and evolutionary processes that sustain it.

Genetic diversity is the “fundamental currency of diversity” (Williams and Humphires,

1996) that is responsible for variation between individuals, populations and species.
Therefore, it is an important aspect of any discussion of biodiversity. The interactions
between the individual organisms (e.g., reproductive behavior, predation, parasitism) of a
population or community, and their specializations for their environment (including ways
in which they might modify the environment itself) are important functional aspects of
biodiversity. These functional aspects can determine the diversity of different
communities and ecosystems.

There is also an important spatial component to biodiversity. The structure of

communities and ecosystems (e.g. the number of individuals and species present) can
vary in different parts of the world. Similarly, the function of these communities and
ecosystems (i.e. the interactions between the organisms present) can vary from one place
to another. Different assemblages of ecosystems can characterize quite diverse
landscapes, covering large areas. These spatial patterns of biodiversity are affected by
climate, geology, and physiography (Redford and Richter, 1999).

The structural, functional, and spatial aspects of biodiversity can vary over time;
therefore there is a temporal component to the analysis of biodiversity. For example,
there can be daily, seasonal, or annual changes in the species and number of organisms
present in an ecosystem and how they interact. Some ecosystems change in size or
structure over time (e.g. forest ecosystems may change in size and structure because of
the effects of natural fires, wetlands gradually silt up and decrease in size). Biodiversity
also changes over a longer-term, evolutionary, time-scale. Geological processes (e.g.,
plate tectonics, orogenesis, erosion), changes in sea-level (marine transgressions and
regressions), and changes in climate cause significant, long-term changes to the structural
and spatial characteristics of global biodiversity. The processes of natural selection and
species evolution, which may often be associated with the geological processes, also
result in changes to local and global flora and fauna.

Many people consider humans to be a part of nature, and therefore a part of biodiversity.
On the other hand, some people (e.g., Redford and Richter, 1999 ) confine biodiversity to
natural variety and variability, excluding biotic patterns and ecosystems that result from
human activity, even though it is difficult to assess the "naturalness" of an ecosystem
because human influence is so pervasive and varied ( Hunter, 1996; Angermeier, 2000;
Sanderson et al.,2002). If one takes humans as part of nature, then cultural diversity of
human populations and the ways that these populations use or otherwise interact with
habitats and other species on Earth are a component of biodiversity too. Other people
make a compromise between totally including or excluding human activities as a part of
biodiversity. These biologists do not accept all aspects of human activity and culture as
part of biodiversity, but they do recognize that the ecological and evolutionary diversity
of domestic species, and the species composition and ecology of agricultural ecosystems
are part of biodiversity. (For further discussion see the modules on Human evolution and
Cultural Diversity; in preparation.)

Glossary
Biodiversity:
the variety of life on Earth at all its levels, from genes to ecosystems, and the
ecological and evolutionary processes that sustain it
Plate Tectonics:
the forces acting on the large, mobile pieces (or "plates") of the Earth's lithosphere
(the upper part of the mantle and crust of the Earth where the rocks are rigid
compared to those deeper below the Earth's surface) and the movement of those
"plates".
Orogenesis:
the process of mountain building.

• U.S. Congress Office of Technology Assessment,

"Technologies to Maintain Biological Diversity," 1987:

"Biological diversity is the variety and variability among

living organisms and the ecological complexes in which
they occur. Diversity can be defined as the number of different
items and their relative frequency. For biological diversity,
these items are organized at many levels, ranging from
complete ecosystems to the chemical structures that are the
molecular basis of heredity. Thus, the term encompasses
different ecosystems, species, genes, and their relative
abundance."

• Jones and Stokes Associates' "Sliding Toward Extinction:

The State of California's Natural Heritage," 1987:

"Natural diversity, as used in this report, is synonymous with

biological diversity... To the scientist, natural diversity has a
variety of meanings. These include: 1) the number of
different native species and individuals in a habitat or
geographical area; 2) the variety of different habitats within
an area; 3) the variety of interactions that occur between
different species in a habitat; and 4) the range of genetic
variation among individuals within a species."
• D.B. Jensen, M. Torn, and J. Harte, "In Our Own Hands: A
Strategy for Conserving Biological Diversity in California,"
1990:

"Biological diversity, simply stated, is the diversity of life...As

defined in the proposed US Congressional Biodiversity
Act, HR1268 (1990), "biological diversity means the full
range of variety and variability within and among living
organisms and the ecological complexes in which they
occur, and encompasses ecosystem or community
diversity, species diversity, and genetic diversity."

Genetic diversity is the combination of different genes found

within a population of a single species, and the pattern of
variation found within different populations of the same
species. Coastal populations of Douglas fir are genetically
different from Sierran populations. Genetic adaptations to local
conditions such as the summer fog along the coast or hot
summer days in the Sierra result in genetic differences
between the two populations of the same species.

Species diversity is the variety and abundance of different

types of organisms which inhabit an area. A ten square mile
area of Modoc County contains different species than does a
similar sized area in San Bernardino County.

Ecosystem diversity encompasses the variety of habitats that

occur within a region, or the mosaic of patches found within a
landscape. A familiar example is the variety of habitats and
environmental parameters that constitute the San Francisco
Bay-Delta ecosystem: grasslands, wetlands, rivers, estuaries,
fresh and salt water."

• Keystone Center, "Final Consensus Report of the Keystone

Policy Dialogue on Biological Diversity on Federal Lands,"
1991:

"In the simplest of terms, biological diversity is the variety of

life and its processes; and it includes the variety of living
organisms, the genetic differences among them, and the
communities and ecosystems in which they occur.

Because biological diversity is so complex, and much of it is

hidden from our view, unknown, or both, it is necessary to
establish means of addressing its distinct and measurable
parts. Most basic of these is genetic variation. Genetic
variation within and between populations of species affects
their physical characteristics, viability, productivity, resilience to
stress, and adaptability to change.

A second, more easily recognized aspect of biological diversity

is distinct species. Some species, such as American elk,
rainbow trout, and ponderosa pine are plentiful. Others such as
the red cockaded woodpecker, Siler's pincushion cactus, or
grizzly bear, have populations that are much reduced or may
even face extinction. Conserving biological diversity includes
perpetuating native species in numbers and distributions that
provide a high likelihood of continued existence.
Associations of species are a third element of biological
diversity. These associations are often called biological
communities, usually recognized as distinct stands, patches, or
sites, such as old-growth forests, riparian areas, or wetlands.
Communities form the biotic parts of ecosystems. The variety
of species in an ecosystem is a function of its structural and
functional characteristics and the diversity of its ecological
processes, and the physical environment.

Finally, at large geographic scales -- from watersheds to the

entire biosphere -- biological diversity includes variety in the
kinds of ecosystems, their patterns, and linkages across
regional landscapes. It is from these large, regional
landscapes, such as the Southern Appalachian Highlands,
Sierra Nevada, and Northern Continental Divide, that people
must derive sustainable yields of resources while perpetuating
multiple intact examples of biologically diverse ecosystems.
This hierarchy of the parts and processes of biological diversity
is admittedly artificial, and it has a distinct human context.
However, it provides a focus for a concept that is infinitely
varied and dynamic and that must be addressed in light of the
full spectrum of human needs and aspirations."

• World Resources Institute, World Conservation Union, and

United Nations Environment Programme, "Global Biodiversity
Strategy," 1992:

"Biodiversity is the totality of genes, species, and

ecosystems in a region... Biodiversity can be divided into
three hierarchical categories -- genes, species, and
ecosystems -- that describe quite different aspects of living
systems and that scientists measure in different ways.

Genetic diversity refers to the variation of genes within

species. This covers distinct populations of the same species
(such as the thousands of traditional rice varieties in India) or
genetic variation within a populations (high among Indian
rhinos, and very low among cheetahs)...

Species diversity refers to the variety of species within a

region. Such diversity can be measured in many ways, and
scientists have not settled on a single best method. The
number of species in a region -- its species "richness" -- is one
often- used measure, but a more precise measurement,
"taxonomic diversity", also considers the relationship of
species to each other. For example, an island with two species
of birds and one species of lizard has a greater taxonomic
diversity than an island with three species of birds but no
lizards...

Ecosystem diversity is harder to measure than species or

genetic diversity because the "boundaries" of communities --
associations of species -- and ecosystems are elusive.
Nevertheless, as long as a consistent set of criteria is used to
define communities and ecosystems, their numbers and
distribution can be measured..."
• Edward Grumbine, "Ghost Bears: Exploring the Biodiversity
Crisis," 1993:

"There is much more to biodiversity than the numbers of

species and kinds of ecosystems. Ecologist Jerry Franklin
portrays ecosystems as having three primary attributes:
composition, structure, and function.

Ecosystem components are the inhabiting species in all their

variety and richness. Many different species, gene-pool
abundance, and unique populations are what most people
think of when they hear the term "biodiversity". But there is
much more to consider.

Ecosystem structure refers to the physical patterns of life

forms from the individual physiognomy of a thick-barked
Douglas-fir to the vertical layers of vegetation from delicate
herbs to tree canopies within a single forest stand. An
ecosystem dominated by old, tall trees has a different structure
than one comprised of short, quaking aspen. And there is more
structure in a multilayered forest (herbs, shrubs, young trees,
canopy trees) than in a single sagebrush grassland, prairie, or
salt marsh...

Ecosystem functions are hard to see in action. "You can't

hug a biogeochemical cycle," says one ecologist. But without
the part of the carbon cycle where small invertebrates, fungi,
and microorganisms work to break down wood fiber, the
downed logs in an ancient forest would never decay. Natural
disturbances also play a role. Wildfires release nutrients to the
soil, weed out weak trees, and reset the successional clock.
The energy of falling water creates spawning beds for salmon
even while it carves a mountain's bones. Plants breathe
oxygen into the atmosphere. Ecological processes create
landscapes and diverse environmental conditions out of life
itself.

Ecosystem components, structures, and functions are all

interdependent. To understand biodiversity, one has to think
like a mountain and consider not only the biotic elements of
plants, animals, and other living beings, but also the patterns
and processes that shape volcanoes and forests."

• Reed Noss, "Indicators for Monitoring Biodiversity: A

Hierarchial Approach," Conservation Biology 4(4):355-364.
1990:

"Biodiversity is not simply the number of genes, species,

ecosystems, or any other group of things in a defined area...A
definition of biodiversity that is altogether simple,
comprehensive, and fully operational (i.e. responsive to real-
life management and regulatory questions) is unlikely to be
found. More useful than a definition, perhaps, would be a
characterization of biodiversity that identifies the major
components at several levels of organization.

...(C)omposition, structure, and function...determine, and in fact

constitute, the biodiversity of an area. Composition has to do
with the identity and variety of elements in a collection, and
 includes species lists and measures of species diversity and
genetic diversity. Structure is the physical organization or
pattern of a system, from habitat complexity as measured
within communities to the pattern of patches and other
elements at a landscape scale. Function involves ecological
and evolutionary processes, including gene flow, disturbances,
and nutrient cycling."

Extinction is a natural event and, from a geological perspective, routine. We now know
that most species that have ever lived have gone extinct. The average rate over the past
200 million years is 1-2 species per million species present per year. The average
duration of a species is 1-10 million years (based on the last 200 million years). There
have also been several episodes of mass extinction, when many taxa representing a wide
array of life forms have gone extinct in the same blink of geological time.

In the modern era, due to human actions, species and ecosystems are threatened with
destruction to an extent rarely seen in Earth history. Probably only during the handful of
mass extinction events have so many species been threatened, in so short a time.

What are these human actions that threaten biodiversity? There are many ways to
conceive of these; let's consider two.

First, we can attribute the loss of species and ecosystems to the accelerating
transformation of the Earth by a growing human population. As the human population
passes the 6 billion mark, we have transformed, degraded or destroyed roughly half of the
word's forests. We appropriate roughly half of the world's net primary productivity for
human use. We appropriate most available fresh water, and we harvest virtually all of the
available productivity of the oceans. It is little wonder that species are disappearing and
ecosystems are being destroyed

Over-hunting
Over-hunting has been a significant cause of the extinction of hundreds of species and the
endangerment of many more, such as whales and many African large mammals. Most
extinctions over the past several hundred years are mainly due to over-harvesting for food,
fashion, and profit. Commercial hunting, both legal and illegal (poaching), is the principal
threat. The snowy egret, passenger pigeon, and heath hen are US examples. At US $16,000 per
pound, and US $40,000 to US $100,000 per horn, it is little wonder that some rhino species are
down to only a few thousand individuals, with only a slim hope of survival in the wild. The
recent expansion of road networks into previously remote tropical forests enables the bushmeat
trade, resulting in what some conservationsist describe as "empty forests" as more and more
wild animals are shot for food.
The pet and decorative plant trade falls within this
commercial hunting category, and includes a mix of legal
and illegal activities. The annual trade is estimated to be at
least US $5 billion, with perhaps one-quarter to one-third
of it illegal. Sport or recreational hunting causes no
endangerment of species where it is well regulated, and
may help to bring back a species from the edge of
extinction. Many wildlife managers view sport hunting as
the principal basis for protection of wildlife.
Nova Scotia Museum of Natural History
(museum.gov.ns.ca/mnh)
While over-hunting, particularly illegal poaching, remains Passenger Pigeon
a serious threat to certain species, for the future, it is
globally less important than other factors mentioned next.

Habitat loss, degradation, fragmentation

Habitat loss, degradation, and fragmentation are important causes of known extinctions. As
deforestation proceeds in tropical forests, this promises to become the main cause of mass
extinctions caused by human activity.

All species have specific food and habitat needs. The more specific these needs and localized
the habitat, the greater the vulnerability of species to loss of habitat to agricultural land,
livestock, roads and cities. In the future, the only species that survive are likely to be those
whose habitats are highly protected, or whose habitat corresponds to the degraded state
associated with human activity (human commensals).

Habitat damage, especially the conversion of forested

land to agriculture (and, often, subsequent abandonment
as marginal land), has a long human history. It began in
China about 4,000 years ago, was largely completed in
Europe by about 400 years ago, and swept across the US Thinking Point
over the past 200 years or so. Viewed in this historical The destruction of tropical
context, we are now mopping up the last forests of the rainforests in economically
Pacific Northwest. developing countries may well
have serious worldwide
In the New World tropics, lowland, seasonal, deciduous ramifications (e.g., loss of
forests began to disappear after 1500 with Spanish and species, global warming). How
might foreign policy in first
Portuguese colonization of the New World. These were
world nations be guided to
the forested regions most easily converted to agriculture, support economic growth in
and with a more welcoming climate. The more rainforest states while
forbidding, tropical humid forests came under attack fostering a sustainable
mainly in twentieth century, under the combined environment?
influences of population growth, inequitable land and
income distribution, and development policies that
targeted rain forests as the new frontier to colonize.
Tropical forests are so important because they harbor at least 50 percent, and perhaps more, of
the world's biodiversity. Direct observations, reinforced by satellite data, document that these
forests are declining. The original extent of tropical rain forests was 15 million square km. Now
there remains about 7.5-8 million square km, so half is gone. The current rate of loss is
estimated at near 2 percent annually (100,000 square km destroyed, another 100,000 square km
degraded). While there is uncertainty regarding the rate of loss, and what it will be in future, the
likelihood is that tropical forests will be reduced to 10-25 percent of their original extent by late
twenty-first century.

Habitat fragmentation is a further aspect of habitat loss

that often goes unrecognized. The forest, meadow, or
other habitat that remains generally is in small, isolated
Discussion bits rather than in large, intact units. Each is a tiny island
{Dis: Deforestation in the that can at best maintain a very small population.
Pacific Northwest is a volatile Environmental fluctuations, disease, and other chance
topic.} factors make such small isolates highly vulnerable to
Discuss arguments for forest extinction. Any species that requires a large home range,
harvesting (e.g., supporting such as a grizzly bear, will not survive if the area is too
the local economy, demand for small. Finally, we know that small land units are
wood and paper, rotational
strongly affected by their surroundings, in terms of
harvest, tradition) and
arguments against doing so
climate, dispersing species, etc. As a consequence, the
(e.g., soil erosion, habitat loss, ecology of a small isolate may differ from that of a
threat to species including similar ecosystem on a larger scale.
salmon). What types of
conciliatory approaches might For the future,
be taken? habitat loss,
degradation, and
fragmentation combined is the single most important Thinking Point
factor in the projected extinction crisis. In Chicago, an infestation of
Asian longhorned beetles in
Invasion of non-native species 1998--introduced inadvertently
Invasion of non-native species is an important and often through packing crates
overlooked cause of extinctions. The African Great imported from China--forced
Lakes--Victoria, Malawi and Tanganyika--are famous the destruction of hundreds of
trees to control its spread.
for their great diversity of endemic species, termed
Create a list of introduced non-
"species flocks," of cichlid fishes. In Lake Victoria, a native species you know of,
single, exotic species, the Nile Perch, has become and how they impacted the
established and may cause the extinction of most of the environment.
native species, by simply eating them all. It was a
purposeful introduction for subsistence and sports
fishing, and a great disaster.

Of all documented extinctions since 1600, introduced species appear to have played a role in at
least half. The clue is the disproportionate number of species lost from islands: some 93 percent
of 30 documented extinctions of species and sub-species of amphibians and reptiles, 93 percent
of 176 species and sub-species of land and freshwater birds, but only
27 percent of 114 species and subspecies of mammals. Why are island
species so vulnerable, and why is this evidence of the role of non-
indigenous species? Islands are laboratories for evolution.

Domino effects
Domino effects occur when the removal of one species (an extinction
event) or the addition of one species (an invasion event) affects the
entire biological system. Domino effects are especially likely when
two or more species are highly interdependent, or when the affected Jil M. Swearingen, US National Park
Service, Washington, D.C.; PCA-
species is a "keystone" species, meaning that it has strong connections APWG
to many other species. Oriental bittersweet

A keystone species is one whose influence on others is disproportionately great. A seminal

study of marine invertebrates in the rocky intertidal region of Washington State found that the
top predator, a starfish, facilitated the coexistence of many other invertebrates by selectively
consuming mussels, which otherwise would crowd out other organisms. Thus a keystone
species is one whose presence or absence both directly and indirectly influences other species
through food web connectivity. Contrary to what some may think, not all species are
"keystones", and it requires careful experimental studies to identify keystone species.

Pollution
Pollution from chemical contaminants certainly poses a further threat to species and ecosystems.
While not commonly a cause of extinction, it likely can be for species whose range is extremely
small, and threatened by contamination. Several species of desert pupfish, occurring in small
isolated pools in the US Southwest, are examples.

Climate change
A changing global climate threatens species and ecosystems. The distribution of species
(biogeography) is largely determined by climate, as is the distribution of ecosystems and plant
vegetation zones (biomes). Climate change may simply shift these distributions but, for a
number of reasons, plants and animals may not be able to adjust. The pace of climate change
almost certainly will be more rapid than most plants are able to migrate.

The presence of roads, cities, and other barriers associated with human
presence may provide no opportunity for distributional shifts. Parks
and nature reserves are fixed locations. The climate that characterizes
present-day Yellowstone Park will shift several hundred miles
northward. The park itself is a fixed location. For these reasons, some
species and ecosystems are likely to be eliminated by climate change.
Mountaintop species are especially vulnerable. The plants and animals
found on high mountains of the American West include many
remnants of a Pleistocene fauna that long ago was displaced toward the
Consider how the
extinction of one species arctic, or upslope. With further warming, many of these mountaintop
might affect other species species likely will be eliminated.
A changing climate will have many other effects. The southern extent of the Everglades, today
the site of the most ambitious and expensive restoration project ever undertaken, may be
underwater, along with significant areas of human habitation. Agricultural production likely
will show regional variation in gains and losses, depending upon crops and climate. Some coral
reefs will expand, and others will contract or die off. Ecological changes due to an altered
climate are difficult to forecast, but expected to be serious.

As a consequence of these multiple forces, many scientists fear that by end of next century,
perhaps 25 percent of existing species will be lost.