What is Biodiversity
Collection Editor:
Nora Bynum
What is Biodiversity
Collection Editor:
Nora Bynum
Authors:
Robert Ahlfinger
James Gibbs
Ian Harrison
Melina Laverty
Eleanor Sterling
Online:
CONNEXIONS
Rice University, Houston, Texas
©2008 Nora Bynum
This selection and arrangement of content is licensed under the Creative Commons Attribution License:
http://creativecommons.org/licenses/by/3.0/
Table of Contents
1 Global Processes ............................................................................ 1
2 Definition of Biodiversity ................................................................... 5
3 Spatial Gradients in Biodiversity ........................................................... 7
4 Introduction to the Biodiversity Hierarchy................................................. 9
5 What is Biodiversity? A comparison of spider communities .............................. 11
6 Species Diversity ........................................................................... 25
7 Alpha, Beta, and Gamma Diversity ....................................................... 31
8 Introduction to Utilitarian Valuation of Biodiversity...................................... 33
9 Biodiversity over Time..................................................................... 35
10 A Brief History of Life on Earth ......................................................... 37
11 Ecosystem Diversity ...................................................................... 39
12 Population Diversity ...................................................................... 41
13 Biogeographic Diversity................................................................... 43
14 Community Diversity ..................................................................... 45
15 Ecoregions................................................................................. 47
16 Extinction.................................................................................49
17 Landscape Diversity ...................................................................... 51
18 Ecological Value........................................................................... 53
Glossary ......................................................................................55
Bibliography..................................................................................59
Index ......................................................................................... 66
Attributions ..................................................................................67
iv
Chapter 1
Global Processes1
1.1 Atmosphere and Climate Regulation
Life on earth plays a critical role in regulating the earth's physical, chemical, and geological properties, from
influencing the chemical composition of the atmosphere to modifying climate.
About 3.5 billion years ago, early life forms (principally cyanobacteria) helped create an oxygenated
atmosphere through photosynthesis, taking up carbon dioxide from the atmosphere and releasing oxygen
(Schopf 1983[87]; Van Valen 1971 [104]). Over time, these organisms altered the composition of the atmo-
sphere, increasing oxygen levels, and paved the way for organisms that use oxygen as an energy source
(aerobic respiration), forming an atmosphere similar to that existing today.
Carbon cycles on the planet between the land, atmosphere, and oceans through a combination of physical,
chemical, geological, and biological processes (IPCC 2001 [73]). One key way biodiversity influences the
composition of the earth's atmosphere is through its role in carbon cycling in the oceans, the largest reservoir
for carbon on the planet (Gruber and Sarmiento[36], in press). In turn, the atmospheric composition of
carbon influences climate. Phytoplankton (or microscopic marine plants) play a central role in regulating
atmospheric chemistry by transforming carbon dioxide into organic matter during photosynthesis. This
carbon-laden organic matter settles either directly or indirectly (after it has been consumed) in the deep
ocean, where it stays for centuries, or even thousands of years, acting as the major reservoir for carbon
on the planet. In addition, carbon also reaches the deep ocean through another biological process - the
formation of calcium carbonate, the primary component of the shells in two groups of marine organisms
coccolithophorids (a phytoplankton) and foraminifera (a single celled, shelled organism that is abundant in
many marine environments). When these organisms die, their shells sink to the bottom or dissolve in the
water column. This movement of carbon through the oceans removes excess carbon from the atmosphere
and regulates the earth's climate.
Over the last century, humans have changed the atmosphere's composition by releasing large amounts of
carbon dioxide. This excess carbon dioxide, along with other 'greenhouse' gases, is believed to be heating
up our atmosphere and changing the world's climate, leading to 'global warming'. There has been much
debate about how natural processes, such as the cycling of carbon through phytoplankton in the oceans, will
respond to these changes. Will phytoplankton productivity increase and thereby absorb the extra carbon
from the atmosphere? Recent studies suggest that natural processes may slow the rate of increase of carbon
dioxide in the atmosphere, but it is doubtful that either the earth's oceans or its forests can absorb the
entirety of the extra carbon released by human activity (Falkowski et al. 2000[25]).
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CHAPTER 1. GLOBAL PROCESSES
1.2 Land Use Change and Climate Regulation
The energy source that ultimately drives the earth's climate is the sun. The amount of solar radiation
absorbed by the earth depends primarily on the characteristics of the surface. Although the link between
solar absorption, thermodynamics, and ultimately climate is very complex, newer studies indicate that
vegetation cover and seasonal variation in vegetation cover affects climate on both global and local scales.
New generations of atmospheric circulation models are increasingly able to incorporate more complex data
related to these parameters (Sellers et al. 1997[90]). Besides regulating the atmosphere's composition, the
extent and distribution of different types of vegetation over the globe modifies climate in three main ways:
* affecting the reflectance of sunlight (radiation balance);
* regulating the release of water vapor (evapotranspiration); and
* changing wind patterns and moisture loss (surface roughness).
The amount of solar radiation reflected by a surface is known as its albedo; surfaces with low albedo reflect
a small amount of sunlight, those with high albedo reflect a large amount. Different types of vegetation have
different albedos; forests typically have low albedo, whereas deserts have high albedo. Deciduous forests are
a good example of the seasonal relationship between vegetation and radiation balance. In the summer, the
leaves in deciduous forests absorb solar radiation through photosynthesis; in winter, after their leaves have
fallen, deciduous forests tend to reflect more radiation. These seasonal changes in vegetation modify climate
in complex ways, by changing evapotranspiration rates and albedo (IPCC 2001 [73]).
Vegetation absorbs water from the soil and releases it back into the atmosphere through evapotranspi-
ration, which is the major pathway by which water moves from the soil to the atmosphere. This release of
water from vegetation cools the air temperature. In the Amazon region, vegetation and climate is tightly
coupled; evapotranspiration of plants is believed to contribute an estimated fifty percent of the annual
rainfall (Salati 1987[85]). Deforestation in this region leads to a complex feedback mechanism, reducing
evapotranspiration rates, which leads to decreased rainfall and increased vulnerability to fire (Laurance and
Williamson 2001 [56]).
Deforestation also influences the climate of cloud forests in the mountains of Costa Rica. The Monteverde
Cloud Forest harbors a rich diversity of organisms, many of which are found nowhere else in the world.
However, deforestation in lower-lying lands, even regions over 50 kilometers way, is changing the local climate,
leaving the "cloud" forest cloudless (Lawton et al. 2001 [57]). As winds pass over deforested lowlands, clouds
are lifted higher, often above the mountaintops, reducing the ability for cloud forests to form. Removing
the clouds from a cloud forest dries the forest, so it can no longer support the same vegetation or provide
appropriate habitat for many of the species originally found there. Similar patterns may be occurring in
other, less studied montane cloud forests around the world.
Different vegetation types and topographies have varying surface roughness, which change the flow
of winds in the lower atmosphere and in turn influences climate. Lower surface roughness also tends to
reduce surface moisture and increase evaporation. Farmers apply this knowledge when they plant trees to
create windbreaks (Johnson et al. 2003[50]). Windbreaks reduce wind speed and change the microclimate,
increase surface roughness, reduce soil erosion, and modify temperature and humidity. For many field crops,
windbreaks increase yields and production efficiency. They also minimize stress on livestock from cold winds.
1.3 Soil and Water Conservation
Biodiversity is also important for global soil and water protection. Terrestrial vegetation in forests and other
upland habitats maintain water quality and quantity, and controls soil erosion.
In watersheds where vegetation has been removed, flooding prevails in the wet season and drought in
the dry season. Soil erosion is also more intense and rapid, causing a double effect: removing nutrient-rich
topsoil and leading to siltation in downstream riverine and ultimately oceanic environments. This siltation
harms riverine and coastal fisheries as well as damaging coral reefs (Turner and Rabalais 1994 [98]; van
Katwijk et al. 1993[103]).
3
One of the most productive ecosystems on earth, wetlands have water present at or near the surface of
the soil or within the root zone, all year or for a period of time during the year, and the vegetation there
is adapted to these conditions. Wetlands are instrumental for the maintenance of clean water and erosion
control. Microbes and plants in wetlands absorb nutrients and in the process filter and purify water of
pollutants before they can enter coastal or other aquatic ecosystems.
Wetlands also reduce flood, wave, and wind damage. They retard the flow of floodwaters and accumulate
sediments that would otherwise be carried downstream or into coastal areas. Wetlands also serve as breeding
grounds and nurseries for fish and support thousands of bird and other animal species.
1.4 Nutrient Cycling
Nutrient cycling is yet another critical service provided by biodiversity - particularly by microorganisms.
Fungi and other microorganisms in soil help break down dead plants and animals, eventually converting this
organic matter into nutrients that enrich the soil (Pimentel et al. 1995[75]).
Nitrogen is essential for plant growth, and an insufficient quantity of it limits plant production in both
natural and agricultural ecosystems. While nitrogen is abundant in the atmosphere, only a few organisms
(commonly known as nitrogen-fixing bacteria) can use it in this form. Nitrogen-fixing bacteria extract
nitrogen from the air, and transform it into ammonia, then other bacteria further break down this ammonia
into nitrogenous compounds that can be absorbed and used by most plants. In addition to their role in
decomposition and hence nutrient cycling, microorganisms also help detoxify waste, changing waste products
into forms less harmful to humans.
1.5 Pollination and Seed Dispersal
An estimated 90 percent of flowering plants depend on pollinators such as wasps, birds, bats, and bees,
to reproduce. Plants and their pollinators are increasingly threatened around the world (Buchmann and
Nabhan 1995[13]; Kremen and Ricketts 2000[54]). Pollination is critical to most major crops and virtually
impossible to replace. For instance, imagine how costly fruit would be (and how little would be available) if
its natural pollinators no longer existed and each developing flower had to be fertilized by hand.
Many animal species are important dispersers of plant seeds. It has been hypothesized that the loss
of a seed disperser could cause a plant to become extinct. At present, there is no example where this has
occurred. A famous example that has often been cited previously is the case of the dodo (Raphus cucullatus)
and the tambalacoque (Sideroxylon grandiflorum). The dodo, a large flightless bird that inhabited the island
of Mauritius in the Indian Ocean, became extinct due to overhunting in the late seventeenth century. It
was once thought that the tambalacoque, a now endangered tree, depended upon the dodo to germinate its
hard-cased seeds (Temple 1977[96]). In the 1970s, only 13 trees remained and it was thought the tree had
not reproduced for 300 years. The seeds of the tree have a very hard coat, as an experiment they were fed
to a turkey; after passing through its gizzard the seeds were viable and germinated. This experiment led
scientists to believe that the extinction of the dodo was coupled to the tambalacoque's inability to reproduce.
However, this hypothesis has not stood up to further scrutiny, as there were several other species (including
three now extinct species, a large-billed parrot, a giant tortoise, and a giant lizard) that were also capable
of cracking the seed (Witmar and Cheke 1991[111]; Catling 2001 [16]). Thus many factors, including the
loss of the dodo, could have contributed to the decline of the tambalacoque. (For further details of causes
of extinction see Historical Perspectives on Extinction and the Current Biodiversity Crisis). Unfortunately,
declines and/or extinctions of species are often unobserved and thus it is difficult to tease out the cause
of the end result, as multiple factors are often operating simultaneously. Similar problems exist today in
understanding current population declines. For example, in a given species, population declines may be
caused by loss of habitat, loss in prey species or loss of predators, a combination of these factors, or possibly
some other yet unidentified cause, such as disease.
In the pine forests of western North America, corvids (including jays, magpies, and crows), squirrels,
4 CHAPTER 1. GLOBAL PROCESSES
and bears play a role in seed dispersal. The Clark's nutcracker (Nucifraga columbiana) is particularly well
adapted to dispersal of whitebark pine (Pinus albicaulis) seeds (Lanner 1996[55]). The nutcracker removes
the wingless seeds from the cones, which otherwise would not open on their own. Nutcrackers hide the
seeds in clumps. When the uneaten seeds eventually grow, they are clustered, accounting for the typical
distribution pattern of whitebark pine in the forest.
In tropical areas, large mammals and frugivorous birds play a key role in dispersing the seeds of trees and
maintaining tree diversity over large areas. For example, three-wattled bellbirds (Procnias tricarunculata)
are important dispersers of tree seeds of members of the Lauraceae family in Costa Rica. Because bellbirds
return again and again to one or more favorite perches, they take the fruit and its seeds away from the
parent tree, spreading Lauraceae trees throughout the forest (Wenny and Levy 1998[107]).
Chapter 2
Definition of Biodiversityl
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 [32]). 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[110]) 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 ecosys-
tems (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 land-
scapes, covering large areas. These spatial patterns of biodiversity are affected by climate, geology, and
physiography (Redford and Richter, 1999[79]).
The structural, functional, and spatial aspects of biodiversity can vary over time; therefore there is a tem-
poral 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 [79]) confine biodiversity to natural variety and variability,
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6 CHAPTER 2. DEFINITION OF BIODIVERSITY
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[40];
Angermeier, 2000[2]; Sanderson et al.,2002[86]). 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.)
Chapter 3
Spatial Gradients in Biodiversityl
Generally speaking, warm tropical ecosystems are richer in species than cold temperate ecosystems at high
latitudes (see Gaston and Williams, 1996[34], for general discussion). A similar pattern is seen for higher
taxonomic groups (genera, families). Various hypotheses (e.g., environmental patchiness, solar energy, pro-
ductivity; see Blackburn and Gaston, 1996[11]) have been raised to explain these patterns. For example, it
is assumed that warm, moist, tropical environments, with long day-lengths provide organisms with more re-
sources for growth and reproduction than harsh environments with low energy resources (Hunter, 2002[41]).
When environmental conditions favor the growth and reproduction of primary producers (e.g., aquatic algae,
corals, terrestrial flora) then these may support large numbers of secondary consumers, such as small her-
bivores, which also support a more numerous and diverse fauna of predators. In contrast, the development
of primary producers in colder temperate ecosystems is constrained by seasonal changes in sunlight and
temperature. Consequently, these ecosystems may support a less diverse biota of secondary consumers and
predators.
Recently, (Allen et al. 2002[1]) developed a model for the effect of ambient temperature on metabolism,
and hence generation time and speciation rates, and used this model to explain the latitudinal gradient in
biodiversity. However, these authors also noted that the principles that underlie these spatial pattern of
biodiversity are still not well understood.
Species and ecosystem diversity is also known to vary with altitude Walter (1985)[105] and Gaston and
Williams (1996: 214-215)[34]. Mountainous environments, also called orobiomes, are subdivided vertically
into altitudinal belts, such as montane, alpine and nival, that have quite different ecosystems. Climatic
conditions at higher elevations (e.g., low temperatures, high aridity) can create environments where relatively
few species can survive. Similarly, in oceans and freshwaters there are usually fewer species as one moves to
increasing depths below the surface. However, in the oceans there may be a rise in species richness close to
the seabed, which is associated with an increase in ecosystem heterogeneity.
By mapping spatial gradients in biodiversity we can also identify areas of special conservation interest.
Conservation biologists are interested in areas that have a high proportion of endemic species, i.e., species
whose distributions are naturally restricted to a limited area. It is obviously important to conserve these
areas because much of their flora and fauna, and therefore the ecosystems so-formed, are found nowhere
else. Areas of high endemism are also often associated with high species richness (see Gaston and Spicer,
1998 [33] for references).
Some conservation biologists have focused their attention on areas that have high levels of endemism (and
hence diversity) that are also experiencing a high rate of loss of ecosystems; these regions are biodiversity
hotspots. Because biodiversity hotspots are characterized by localized concentrations of biodiversity under
threat, they represent priorities for conservation action (Sechrest et al., 2002[89]). A terrestrial biodiver-
sity hotspot is defined quantitatively as an area that has at least 0.5%, or 1,500 of the world's ca. 300,000
species of green plants (Viridiplantae), and that has lost at least 70% of its primary vegetation (Myers et
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8 CHAPTER 3. SPATIAL GRADIENTS IN BIODIVERSITY
al., 2000[70]; Conservation International, 2002[46]). Marine biodiversity hotspots are quantitatively
defined based on measurements of relative endemism of multiple taxa (species of corals, snails, lobsters,
fishes) within a region and the relative level of threat to that region (Roberts et al., 2002[81]). According
to this approach, the Philippine archipelago and the islands of Bioko, Sao Tome, Principe and Annobon in
the eastern Atlantic Gulf of Guinea are ranked as two of the most threatened marine biodiversity hotspot
regions.
Conservation biologists may also be interested in biodiversity coldspots; these are areas that have
relatively low biological diversity but also include threatened ecosystems (Kareiva and Marvier, 2003[51]).
Although a biodiversity coldspot is low in species richness, it can also be important to conserve, as it may be
the only location where a rare species is found. Extreme physical environments (low or high temperatures
or pressures, or unusual chemical composition) inhabited by just one or two specially adapted species are
coldspots that warrant conservation because they represent unique environments that are biologically and
physically interesting. For further discussion on spatial gradients in biodiversity and associated conservation
practices see the related modules on "Where is the world's biodiversity?" and "Conservation Planning at a
Regional Scale."
Chapter 4
Introduction to the Biodiversity
Hierarchyl
To effectively conserve biodiversity, we need to be able to define what we want to conserve, determine where
it currently occurs, identify strategies to help conserve it, and track over time whether or not these strategies
are working. The first of these items, defining what we want to conserve, is complicated by the remarkable
diversity of the organisms themselves. This is a product of the genetic diversity of the organisms, that is,
variation in the DNA (deoxyribonucleic acid) that makes up the genes of the organisms.
Genetic diversity among organisms exists at the following different levels:
" within a single individual;
" between different individuals of a single population;
" between different populations of a single species (population diversity);
" between different species (species diversity).
It can be difficult, in some cases, to establish the boundaries between these levels of diversity. For
example, it may be difficult to interpret whether variation between groups of individuals represents diversity
between different species, or represents diversity only between different populations of the same species.
Nevertheless, in general terms, these levels of genetic diversity form a convenient hierarchy for describing
the overall diversity of organisms on Earth.
Similarly, the functional and spatial aspects of biodiversity can also be discussed at a number of different
levels; for example, diversity within or between communities, ecosystems, landscapes, biogeographical
regions, and ecoregions.
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10 CHAPTER 4. INTRODUCTION TO THE BIODIVERSITY HIERARCHY
Chapter 5
What is Biodiversity? A comparison of
spider communities
5.1 Objectives
To explore through classification of life forms the concept of biological diversity as it occurs at various
taxonomic levels.
5.2 Procedures
Spiders are a highly species rich group of invertebrates that exploit a wide variety of niches in virtually
all the earth's biomes. Some species of spiders build elaborate webs that passively trap their prey whereas
others are active predators that ambush or pursue their prey. Given spiders' taxonomic diversity as well as
the variety of ecological niches breadth along with the ease of catching them, spiders can represent useful,
fairly easily measured indicators of environmental change and community level diversity.
This exercise focuses on classifying and analyzing spider communities to explore the concept of biological
diversity and experience its application to decision making in biological conservation. The exercise can be
undertaken in three parts, depending on your interest level.
" Level (1) - You will gain experience in classifying organisms by sorting a hypothetical collection of
spiders from a forest patch and determining if the spider collection is adequate to accurately represent
the overall diversity of spiders present in the forest patch.
" Level (2) - If you wish to explore further, you can sort spider collections made at four other forest
patches in the same region and contrast spider communities in terms of their species richness, species
diversity, and community similarity. You will apply this information to make decisions about the
priority that should be given to protecting each forest patch in order to conserve the regional pool of
spider diversity.
" Level (3) - If you wish to explore the concepts of biodiversity yet further, you will next take into
account the evolutionary relationships among the families of spiders collected. This phylogenetic
perspective will augment your decision making about priorities for patch protection by accounting
for evolutionary distinctiveness in addition to diversity and distinctiveness at the community level.
Once you have worked through these concepts and analyses you will have a much enhanced familiarity with
the subtleties of what biological diversity is.
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12 CHAPTER 5. WHAT IS BIODIVERSITY? A COMPARISON OF SPIDER
COMMUNITIES
5.3 Level 1: Sorting and Classifying a Spider Collection and Assess-
ing its Comprehensiveness
Obtain a paper copy of the spider collection for forest patch "1." The spiders were captured by a biologist
traveling along transects through the patch and striking a random series of 100 tree branches. All spiders
dislodged that fell onto an outstretched sheet were collected and preserved in alcohol. They have since been
spread out on a tray for you to examine. The spider collection is hypothetical but the species pictured are
actual spiders that occur in central Africa (illustrations used are from Berland 1955[9]).
The next task is for you to sort and identify the spiders. To do this you have to identify all the specimens
in the collection. To classify the spiders look for external characters that all members of a particular group of
spiders have in common but that are not shared by other groups of spiders. For example, leg length, hairiness,
relative size of body segments, or abdomen patterning and abdomen shape all might be useful characters.
Look for groups of morphologically indistinguishable spiders, and describe briefly the set of characters unique
to each group. These operational taxonomic units that you define will be considered separate species. To
assist you in classifying these organisms, a diagram of key external morphological characters of beetles is
provided (Figure 5.1). Note that most spider identification depends on close examination of spider genitalia.
For this exercise, however, we will be examining gross external characteristics of morphologically dissimilar
species.
13
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Figure 5.1: Basic external characteristics of spiders useful for identifying individuals to species.
Assign each species a working name, preferably something descriptive. For example, you might call a
particular species "spotted abdomen, very hairy" or "short legs, spiky abdomen" Just remember that the
more useful names will be those that signify to you something unique about the species. Construct a table
listing each species, its distinguishing characteristics, the name you have applied to it, and the number of
occurrences of the species in the collection (Figure 5.2).
14 CHAPTER 5. WHAT IS BIODIVERSITY? A COMPARISON OF SPIDER
COMMUNITIES
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Last, ask whether this collection adequately represents the true diversity of spiders in the forest patch
at the time of collection. Were most of the species present sampled or were many likely missed? This is
always an important question to ask to ensure that the sample was adequate and hence can be legitimately
contrasted among sites to, for example, assign areas as low versus high diversity sites.
To do this you will perform a simple but informative analysis that is standard practice for conservation
biologists who do biodiversity surveys. This analysis involves constructing a so-called collector's curve
(Colwell and Coddington 1994[18]). These plot the cumulative number of species observed (y-axis) against
the cumulative number of individuals classified (x-axis). The collector's curve is an increasing function with
a slope that will decrease as more individuals are classified and as fewer species remain to be identified
(Figure 5.3). If sampling stops while the collector's curve is still rapidly increasing, sampling is incomplete
and many species likely remain undetected. Alternatively, if the slope of the collector's curve reaches zero
(flattens out), sampling is likely more than adequate as few to no new species remain undetected.
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individuals classified. The cumulative number of taxa sampled refers to the number of new species
detected.
To construct the collector's curve for this spider collection, choose a specimen within the collection at
random. This will be your first data point, such that X = 1 and Y = 1 because after examining the
first individual you have also identified one new species! Next move consistently in any direction to a new
specimen and record whether it is a member of a new species. In this next step, X = 2, but Y may remain
as 1 if the next individual is not of a new species or it may change to 2 if the individual represents a new
species different from individual 1. Repeat this process until you have proceeded through all 50 specimens
and construct the collector's curve from the data obtained (just plot Y versus X). Does the curve flatten
out? If so, after how many individual spiders have been collected? If not, is the curve still increasing? What
16 CHAPTER 5. WHAT IS BIODIVERSITY? A COMPARISON OF SPIDER
COMMUNITIES
can you conclude from the shape of your collector's curve as to whether the sample of spiders is an adequate
characterization of spider diversity at the site?
5.4 Level 2: Contrasting spider diversity among sites to provide a
basis for prioritizing conservation efforts
In this part of the exercise you are provided with spider collections from 4 other forest patches. The forest
patches have resulted from fragmentation of a once much larger, continuous forest. You will use the spider
diversity information to prioritize efforts for the five different forest patches (including the data from the
first patch which you have already classified). Here are the additional spider collections: (See Figure 5.4,
Figure 5.5, Figure 5.6, and Figure 5.7)
17
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18 CHAPTER 5. WHAT IS BIODIVERSITY? A COMPARISON OF SPIDER
COMMUNITIES
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Figure 5.6
20 CHAPTER 5. WHAT IS
BIODIVERSITY? A COMPARISON OF SPIDER
COMMUNITIES
Site
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25
26
CHAPTER 6. SPECIES DIVERSITY
To count the number of species, we must define what constitutes a species. There are several competing
theories, or "species concepts" (Mayden, 1997[62]). The most widely accepted are the morphological species
concept, the biological species concept, and the phylogenetic species concept.
Although the morphological species concept (MSC) is largely outdated as a theoretical definition, it
is still widely used. According to this concept:
species are the smallest groups that are consistently and persistently distinct, and distinguishable
by ordinary means. (- Cronquist, 1978[?]).
In other words, morphological species concept states that "a species is a community, or a number of re-
lated communities, whose distinctive morphological characters are, in the opinion of a competent systematist,
sufficiently definite to entitle it, or them, to a specific name" (Regan, 1926: 75[80]).
The biological species concept (BSC), as described by Mayr and Ashlock (1991)[64], states that
"a species is a group of interbreeding natural populations that is reproductively isolated from
other such groups".
According to the phylogenetic species concept (PSC), as defined by Cracraft (1983)[19], a species
"is the smallest diagnosable cluster of individual organism [that is, the cluster of organisms are
identifiably distinct from other clusters] within which there is a parental pattern of ancestry and
descent".
These concepts are not congruent, and considerable debate exists about the advantages and disadvantages
of all existing species concepts (for further discussion, see the module on Macroevolution: essentials of
systematics and taxonomy).
In practice, systematists usually group specimens together according to shared features (genetic, mor-
phological, physiological). When two or more groups show different sets of shared characters, and the shared
characters for each group allow all the members of that group to be distinguished relatively easily and consis-
tently from the members of another group, then the groups are considered different species. This approach
relies on the objectivity of the phylogenetic species concept (i.e., the use of intrinsic, shared, characters to
define or diagnose a species) and applies it to the practicality of the morphological species concept, in terms
of sorting specimens into groups (Kottelat, 1995[52], 1997[53]).
Despite their differences, all species concepts are based on the understanding that there are parameters
that make a species a discrete and identifiable evolutionary entity. If populations of a species become
isolated, either through differences in their distribution (i.e., geographic isolation) or through differences in
their reproductive biology (i.e., reproductive isolation), they can diverge, ultimately resulting in speciation.
During this process, we expect to see distinct populations representing incipient species - species in
the process of formation. Some researchers may describe these as subspecies or some other sub-category,
according to the species concept used by these researchers. However, it is very difficult to decide when a
population is sufficiently different from other populations to merit its ranking as a subspecies. For these
reasons, subspecific and infrasubspecific ranks may become extremely subjective decisions of the degree of
distinctiveness between groups of organisms (Kottelat, 1997[53]).
An evolutionary significant unit (ESU) is defined, in conservation biology, as a group of organisms
that has undergone significant genetic divergence from other groups of the same species. According to Ryder,
1986[84] identification of ESUs requires the use of natural history information, range and distribution data,
and results from analyses of morphometrics, cytogenetics, allozymes and nuclear and mitochondrial DNA.
In practice, many ESUs are based on only a subset of these data sources. Nevertheless, it is necessary
to compare data from different sources (e.g., analyses of distribution, morphometrics, and DNA) when
establishing the status of ESUs. If the ESUs are based on populations that are sympatric or parapatric
then it is particularly important to give evidence of significant genetic distance between those populations.
ESUs are important for conservation management because they can be used to identify discrete compo-
nents of the evolutionary legacy of a species that warrant conservation action. Nevertheless, in evolutionary
terms and hence in many systematic studies, species are recognized as the minimum identifiable unit of
27
biodiversity above the level of a single organism (Kottelat, 1997[53]). Thus there is generally more system-
atic information available for species diversity than for subspecific categories and for ESUs. Consequently,
estimates of species diversity are used more frequently as the standard measure of overall biodiversity of a
region.
6.1 Species Diversity as a Surrogate for Global Biodiversity
Global biodiversity is frequently expressed as the total number of species currently living on Earth, i.e.,
its species richness. Between about 1.5 and 1.75 million species have been discovered and scientifically
described thus far (LeCointre and Guyader, 2001[58]; Cracraft, 2002[20]). Estimates for the number of
scientifically valid species vary partly because of differing opinions on the definition of a species.For example,
the phylogenetic species concept recognizes more species than the biological species concept. Also, some
scientific descriptions of species appear in old, obscure, or poorly circulated publications. In these cases,
scientists may accidentally overlook certain species when preparing inventories of biota, causing them to
describe and name an already known species.
More significantly, some species are very difficult to identify. For example, taxonomically "cryptic species"
look very similar to other species and may be misidentified (and hence overlooked as being a different species).
Thus, several different, but similar-looking species, identified as a single species by one scientist, are identified
as completely different species by another scientist. For further discussion of cryptic species, with specific
examples of cryptic frogs from Vietnam, see Inger (1999) [45] and Bain et al., (in press)[7].
Scientists expect that the scientifically described species represent only a small fraction of the total number
of species on Earth today. Many additional species have yet to be discovered, or are known to scientists
but have not been formally described. Scientists estimate that the total number of species on Earth could
range from about 3.6 million up to 117.7 million, with 13 to 20 million being the most frequently cited range
(Hammond, 1995[37]; Cracraft, 2002[20]).
The estimation of total number of species is based on extrapolations from what we already know about
certain groups of species. For example, we can extrapolate using the ratio of scientifically described species to
undescribed species of a particular group of organisms collected from a prescribed area. However, we know so
little about some groups of organisms, such as bacteria and some types of fungi, that we do not have suitable
baseline data from which we can extrapolate our estimated total number of species on Earth. Additionally,
some groups of organisms have not been comprehensively collected from areas where their species richness
is likely to be richest (for example, insects in tropical rainforests). These factors, and the fact that different
people have used different techniques and data sets to extrapolate the total number of species, explain the
large range between the lower and upper figures of 3.6 million and 117.7 million, respectively.
While it is important to know the total number of species of Earth, it is also informative to have some
measure of the proportional representation of different groups of related species (e.g. bacteria, flowering
plants, insects, birds, mammals). This is usually referred to as the taxonomic or phylogenetic diversity.
Species are grouped together according to shared characteristics (genetic, anatomical, biochemical, phys-
iological, or behavioral) and this gives us a classification of the species based on their phylogenetic, or
apparent evolutionary relationships. We can then use this information to assess the proportion of related
species among the total number of species on Earth. Table 6.1: Estimated Numbers of Described Species,
Based on Lecointre and Guyader (2001) contains a selection of well-known taxa.
28
CHAPTER 6. SPECIES DIVERSITY
Estimated Numbers of Described Species, Based on Lecointre and Guyader (2001)
Taxon Taxon Common Number of species N as percentage of
Name described* total number of de-
scribed species*
Bacteria true bacteria 9021 0.5
Archaea archaebacteria 259 0.01
Bryophyta mosses 15000 0.9
Lycopodiophyta clubmosses 1275 0.07
Filicophyta ferns 9500 0.5
Coniferophyta conifers 601 0.03
Magnoliophyta flowering plants 233885 13.4
Fungi fungi 100800 5.8
"Porifera" sponges 10000 0.6
Cnidaria cnidarians 9000 0.5
Rotifera rotifers 1800 0.1
Platyhelminthes flatworms 13780 0.8
Mollusca mollusks 117495 6.7
Annelida annelid worms 14360 0.8
Nematoda nematode worms 20000 1.1
Arachnida arachnids 74445 4.3
Crustacea crustaceans 38839 2.2
Insecta insects 827875 47.4
Echinodermata echinoderms 6000 0.3
Chondrichthyes cartilaginous fishes 846 0.05
Actinopterygii ray-finned bony fishes 23712 1.4
Lissamphibia living amphibians 4975 0.3
Mammalia mammals 4496 0.3
Chelonia living turtles 290 0.02
Squamata lizards and snakes 6850 0.4
continued on next page
29
Aves birds 9672 0.6
Other 193075 11.0
Table 6.1: * The total number of described species is assumed to be 1,747,851. This figure, and the
numbers of species for taxa are taken from LeCointre and Guyader (2001)[58].
Most public attention is focused on the biology and ecology of large, charismatic species such as mammals,
birds, and certain species of trees (e.g., mahogany, sequoia). However, the greater part of Earth's species
diversity is found in other, generally overlooked groups, such as mollusks, insects, and groups of flowering
plants.
30 CHAPTER 6. SPECIES DIVERSITY
Chapter 7
Alpha, Beta, and Gamma Diversityl
Whittaker (1972)[108] described three terms for measuring biodiversity over spatial scales: alpha, beta, and
gamma diversity. Alpha diversity refers to the diversity within a particular area or ecosystem, and is
usually expressed by the number of species (i.e., species richness) in that ecosystem. For example, if we
are monitoring the effect that British farming practices have on the diversity of native birds in a particular
region of the country, then we might want to compare species diversity within different ecosystems, such as
an undisturbed deciduous wood, a well-established hedgerow bordering a small pasture, and a large arable
field. We can walk a transect in each of these three ecosystems and count the number of species we see;
this gives us the alpha diversity for each ecosystem; see Table 7.1: Alpha, beta and gamma diversity for
hypothetical species of birds in three different ecosystems (this example is based on the hypothetical example
given by Meffe et al., 2002; Table 6.1[66]).
If we examine the change in species diversity between these ecosystems then we are measuring the
beta diversity. We are counting the total number of species that are unique to each of the ecosystems
being compared. For example, the beta diversity between the woodland and the hedgerow habitats is 7
(representing the 5 species found in the woodland but not the hedgerow, plus the 2 species found in the
hedgerow but not the woodland). Thus, beta diversity allows us to compare diversity between ecosystems.
Gamma diversity is a measure of the overall diversity for the different ecosystems within a region.
Hunter (2002: 448) ([42]) defines gamma diversity as "geographic-scale species diversity". In the example in
Table 7.1: Alpha, beta and gamma diversity for hypothetical species of birds in three different ecosystems,
the total number of species for the three ecosystems 14, which represent the gamma diversity.
Alpha, beta and gamma diversity for hypothetical species of birds in three different
ecosystems
Hypothetical species Woodland habitat Hedgerow habitat Open field habitat
A X
B X
C X
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32
CHAPTER 7. ALPHA, BETA, AND GAMMA DIVERSITY
D X
E X
F X X
G X X
H X X
X X
x x
K X
L X X
M X
N X
Alpha diversity 10 7 3
Beta diversity Woodland vs. Hedgerow vs. open Woodland vs. open
hedgerow: 7 field: 8 field: 13
Gamma diversity 14
Table 7.1
Chapter 8
Introduction to Utilitarian Valuation of
Biodiversi
Determining the value or worth of biodiversity is complex. Economists typically subdivide utilitarian or
use values of biodiversity into direct use value for those goods that are consumed directly, such as food
or timber, and indirect use value for those services that support the items that are consumed, including
ecosystem functions like nutrient cycling.
There are several less tangible values that are sometimes called non-use or passive values, for things
that we don't use but would consider as a loss if they were to disappear; these include existence value, the
value of knowing something exists even if you will never use it or see it, and bequest value, the value of
knowing something will be there for future generations (Moran and Pearce 1994 [69]). Potential or Option
value refers to the use that something may have in the future; sometimes this is included as a use value,
we have chosen to include it within the passive values here based on its abstract nature. The components
included within the category of "utilitarian" values vary somewhat in the literature. For example, some
authors classify spiritual, cultural, and aesthetic values as indirect use values, whiles others consider them
to be non-use values, differentiated from indirect use values - such as nutrient cycling - because spiritual,
cultural, and aesthetic values for biodiversity are not essential to human survival. Still others consider these
values as separate categories entirely. (See also, Callicott 1997[15], Hunter 2002[44], Moran and Pearce
1994[69], Perlman and Adelson 1997[74], Primack 2002[77], Van Dyke 2003[102]). In this module, we
include spiritual, cultural and aesthetic values as a subset of indirect values or services, as they provide a
service by enriching our lives (Table 8.1: Categories of Values of Biodiversity).
Categories of Values of Biodiversity
Direct Use Value Indirect Use Value Non-Use Values
(Goods) (Services)
Food, medicine, build- Atmospheric and cli- Potential (or Option) Future value either as a
ing material, fiber, fuel mate regulation, polli- Value good or a service
nation, nutrient recy-
cling
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34
CHAPTER 8. INTRODUCTION TO UTILITARIAN VALUATION OF
BIODIVERSITY
Cultural, Spiritual, and Existence Value Value of knowing some-
Aesthetic thing exists
Bequest Value Value of knowing that
something will be there
for future generations
Table 8.1
NOTE: Some authors choose to differentiate Cultural, Spiritual, Aesthetic, and Non-Use Values
from those services that provide basic survival needs such as the air we breathe.
Chapter 9
Biodiversity over Time
The history of life on Earth is described in various publications and web sites (e.g., Speer, B.R. and
A.G. Collins. 2000[92]; Tudge, 2000[97]; Lecointre and Guyader, 2001[59]; Maddison, 2001 [60] Eldredge,
2002[24]); it is also discussed in the module on Macroevolution: essentials of systematics and taxonomy. For
the current purpose of understanding what is biodiversity, it is only necessary to note that that the diversity
of species, ecosystems and landscapes that surround us today are the product of perhaps 3.7 billion (i.e.,
3.7 x 109) to 3.85 billion years of evolution of life on Earth (Mojzsis et al., 1996[67]; Fedo and Whitehouse,
2002[26]).
Thus, the evolutionary history of Earth has physically and biologically shaped our contemporary environ-
ment. As noted in the section on Biogeography (Chapter 13), plate tectonics and the evolution of continents
and ocean basins have been instrumental in directing the evolution and distribution of the Earth's biota.
However, the physical environment has also been extensively modified by these biota. Many existing land-
scapes are based on the remains of earlier life forms. For example, some existing large rock formations are
the remains of ancient reefs formed 360 to 440 million years ago by communities of algae and invertebrates
(Veron, 2000[47]). Very old communities of subterranean bacteria may have been responsible for shaping
many geological processes during the history of the Earth, such as the conversion of minerals from one form
to another, and the erosion of rocks (Fredrickson and Onstott, 1996[30]). The evolution of photosynthetic
bacteria, sometime between 3.5 and 2.75 million years ago Schopf, 1993[88]; Brasier et al., 2002[12]; Hayes,
2002[38]), played an important role in the evolution of the Earth's atmosphere. These bacteria released oxy-
gen into the atmosphere, changing it's composition from the former composition of mainly carbon dioxide,
with other gases such as nitrogen, carbon monoxide, methane, hydrogen and sulphur gases present in smaller
quantities. It probably took over 2 billion years for the oxygen concentration to reach the level it is today
(Hayes, 2002[38]), but the process of oxygenation of the atmosphere led to important evolutionary changes
in organisms so that they could utilize oxygen for metabolism. The rise of animal and plant life on land was
associated with the development of an oxygen rich atmosphere.
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36 CHAPTER 9. BIODIVERSITY OVER TIME
Chapter 10
A Brief History of Life on Earth'
The diversity of species, ecosystems and landscapes that surround us today are the product of perhaps 3.7
billion (i.e., 3.7 x 109) to 3.85 billion years of evolution of life on Earth (Mojzsis et al., 1996[68]; Fedo and
Whitehouse, 2002[?]). Life may have first evolved under harsh conditions, perhaps comparable to the deep-
sea thermal vents where chemo-autotrophic bacteria are currently found (these are organisms that obtain
their energy only from inorganic, chemical sources).
A subterranean evolution of life has also been suggested. Rock layers deep below the continents and
ocean floors, that were previously thought to be too poor in nutrients to sustain life, have now been found
to support thousands of strains of microorganisms. Types of bacteria have been collected from rock samples
almost 2 miles below the surface, at temperatures up to 75 degrees Celsius. These chemo-autotrophic
microorganisms derive their nutrients from chemicals such as carbon, hydrogen, iron and sulphur. Deep
subterranean communities could have evolved underground or originated on the surface and become buried
or otherwise transported down into subsurface rock strata, where they have subsequently evolved in isolation.
Either way, these appear to be very old communities, and it is possible that these subterranean bacteria
may have been responsible for shaping many geological processes during the history of the Earth (e.g.,
the conversion of minerals from one form to another, and the erosion of rocks) (Fredrickson and Onstott,
1996[31]).
The earliest evidence for photosynthetic bacteria - suspected to be cyanobacteria - is dated at sometime
between 3.5 and 2.75 billion years ago (Schopf, 1993[?]; Brasier et al., 2002[?]; Hayes, 2002[?]). These first
photosynthetic organisms would have been responsible for releasing oxygen into the atmosphere. (Photo-
synthesis is the formation of carbohydrates from carbon dioxide and water, through the action of light
energy on a light-sensitive pigment, such as chlorophyll, and usually resulting in the production of oxygen).
Prior to this, the atmosphere was mainly composed of carbon dioxide, with other gases such as nitrogen,
carbon monoxide, methane, hydrogen and sulphur gases present in smaller quantities.
It probably took over 2 billion years, from the initial advent of photosynthesis for the oxygen concentration
in the atmosphere to reach the level it is at today (Hayes, 2002[?]). As oxygen levels rose, some of the
early anaerobic species probably became extinct, and others probably became restricted to habitats that
remained free of oxygen. Some assumed a lifestyle permanently lodged inside aerobic cells. The anaerobic
cells might, initially, have been incorporated into the aerobic cells after those aerobes had engulfed them
as food. Alternatively, the anaerobes might have invaded the aerobic hosts and become parasites within
them. Either way, a more intimate symbiotic relationship subsequently evolved between these aerobic and
anaerobic cells. In these cases the survival of each cell was dependent on the function of the other cell.
The evolution of this symbiotic relationship was an extremely important step in the evolution of more
complex cells that have a nucleus, which is a characteristic of the Eucarya or eucaryotes (eu good, or true;
and karyon kernel, or nucleus). Recent studies of rocks from Western Australia have suggested that the
earliest forms of single-celled eucaryotes might be at least 2.7 billion years old (Anon, 2001[3]). According
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CHAPTER 10. A BRIEF HISTORY OF LIFE ON EARTH
to contemporary theories, there has been sufficient time, over those 2.7 billion years, for some of the genes
of the invading anaerobe to have been lost, or even transferred to the nucleus of the host aerobe cell. As a
result, the genomes of the ancestral invader and ancestral host have become mingled and the two entities
can now be considered as one from a genetic standpoint.
The evolutionary history of the Eucarya is described in various standard references and so is not covered
in detail here. Briefly, eucaryotes constitute three well known groups - the Viridiplantae or green plants,
the Fungi, and the Metazoa or animals. There are also many basal groups of eucaryotes that are extremely
diverse - and many of which are evolutionarily ancient. For example, the Rhodophyta, or red algae, which
might be the sister-group to the Viridiplantae, includes fossil representatives dating from the Precambrian,
1025 billion years ago. The Stramenopiles includes small, single-celled organisms such as diatoms, fungus-
like species of water moulds and downy mildews, and extremely large, multicellular brown seaweeds such as
kelps.
The earliest known green plants are green algae, dating from the Cambrian, at least 500 million years
ago. By the end of the Devonian, 360 million years ago, plants had become quite diverse and included
representatives similar to modern plants. Green plants have been extremely important in shaping the
environment. Fueled by sunlight, they are the primary producers of carbohydrates, sugars that are essential
food resources for herbivores that are then prey to predatory carnivores. The evolution and ecology of
pollinating insects is closely associated with the evolution of the Angiosperms, or flowering plants, since the
Jurassic and Cretaceous periods.
Fungi, which date back to the Precambrian times about 650 to 540 million years ago, are also important
in shaping and sustaining biodiversity. By breaking down dead organic material and using this for their
growth, they recycle nutrients back through ecosystems. Fungi are also responsible for causing several plant
and animal diseases. Fungi also form symbiotic relationships with tree species, often in nutrient-poor soils
such as are found in the humid tropics, allowing their symbiont trees the ability to flourish in what would
otherwise be a difficult environment.
Metazoa, which date to over 500 million years ago have also been responsible for shaping many ecosystems,
from the specialized tubeworms of deep sea, hydrothermal vent communities of the ocean floor, to the birds
living in the high altitudes of the Himalayas, such as the impeyan pheasant and Tibetan snow cock. Many
species of animals are parasitic on other species and can significantly affect the behavior and life-cycles of
their hosts.
Thus, the evolutionary history of Earth has physically and biologically shaped our contemporary environ-
ment. Many existing landscapes are based on the remains of earlier life forms. For example, some existing
large rock formations are the remains of ancient reefs formed 360 to 440 million years ago by communities
of algae and invertebrates (Veron, 2000[48]).
Chapter 11
Ecosystem Diversity
An ecosystem is a community plus the physical environment that it occupies at a given time. An ecosystem
can exist at any scale, for example, from the size of a small tide pool up to the size of the entire biosphere.
However, lakes, marshes, and forest stands represent more typical examples of the areas that are compared
in discussions of ecosystem diversity.
Broadly speaking, the diversity of an ecosystem is dependent on the physical characteristics of the envi-
ronment, the diversity of species present, and the interactions that the species have with each other and with
the environment. Therefore, the functional complexity of an ecosystem can be expected to increase with the
number and taxonomic diversity of the species present, and the vertical and horizontal complexity of the
physical environment. However, one should note that some ecosystems (such as submarine black smokers,
or hot springs) that do not appear to be physically complex, and that are not especially rich in species,
may be considered to be functionally complex. This is because they include species that have remarkable
biochemical specializations for surviving in the harsh environment and obtaining their energy from inorganic
chemical sources (e.g., see discussions of Rothschild and Mancinelli, 2001 [83]).
The physical characteristics of an environment that affect ecosystem diversity are themselves quite com-
plex (as previously noted for community diversity (Chapter 14)). These characteristics include, for example,
the temperature, precipitation, and topography of the ecosystem. Therefore, there is a general trend for
warm tropical ecosystems to be richer in species than cold temperate ecosystems (see "Spatial gradients in
biodiversity (Chapter 3)"). Also, the energy flux in the environment can significantly affect the ecosystem.
An exposed coastline with high wave energy will have a considerably different type of ecosystem than a
low-energy environment such as a sheltered salt marsh. Similarly, an exposed hilltop or mountainside is
likely to have stunted vegetation and low species diversity compared to more prolific vegetation and high
species diversity in sheltered valleys (see Walter, 1985[106], and Smith, 1990[91] for general discussions on
factors affecting ecosystems, and comparative ecosystem ecology).
Environmental disturbance on a variety of temporal and spatial scales can affect the species richness
and, consequently, the diversity of an ecosystem. For example, river systems in the North Island of New
Zealand have been affected by volcanic disturbance several times over the last 25,000 years. Ash-laden floods
running down the rivers would have extirpated most of the fish fauna in the rivers, and recolonization has
been possible only by a limited number of diadromous species (i.e., species, like eels and salmons, that
migrate between freshwater and seawater at fixed times during their life cycle). Once the disturbed rivers
had recovered, the diadromous species would have been able to recolonize the rivers by dispersal through
the sea from other unaffected rivers (McDowall, 1996[65]).
Nevertheless, moderate levels of occasional disturbance can also increase the species richness of an ecosys-
tem by creating spatial heterogeneity in the ecosystem, and also by preventing certain species from dominat-
ing the ecosystem. (See the module on Organizing Principles of the Natural World for further discussion).
Ecosystems may be classified according to the dominant type of environment, or dominant type of
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40 CHAPTER 11. ECOSYSTEM DIVERSITY
species present; for example, a salt marsh ecosystem, a rocky shore intertidal ecosystem, a mangrove swamp
ecosystem. Because temperature is an important aspect in shaping ecosystem diversity, it is also used in
ecosystem classification (e.g., cold winter deserts, versus warm deserts) (Udvardy, 1975[99]).
While the physical characteristics of an area will significantly influence the diversity of the species within
a community, the organisms can also modify the physical characteristics of the ecosystem. For example, stony
corals (Scleractinia) are responsible for building the extensive calcareous structures that are the basis for coral
reef ecosystems that can extend thousands of kilometers (e.g. Great Barrier Reef). There are less extensive
ways in which organisms can modify their ecosystems. For example, trees can modify the microclimate and
the structure and chemical composition of the soil around them. For discussion of the geomorphic influences of
various invertebrates and vertebrates see (Butler, 1995[14]) and, for further discussion of ecosystem diversity
see the module on Processes and functions of ecological systems .
Chapter 12
Population Diversityl
A population is a group of individuals of the same species that share aspects of their genetics or demogra-
phy more closely with each other than with other groups of individuals of that species (where demography
is the statistical characteristic of the population such as size, density, birth and death rates, distribution,
and movement of migration).
Population diversity may be measured in terms of the variation in genetic and morphological features
that define the different populations. The diversity may also be measured in terms of the populations'
demographics, such as numbers of individuals present, and the proportional representation of different age
classes and sexes. However, it can be difficult to measure demography and genetics (e.g., allele frequencies)
for all species. Therefore, a more practical way of defining a population, and measuring its diversity, is by
the space it occupies. Accordingly, a population is a group of individuals of the same species occupying a
defined area at the same time (Hunter, 2002: 144[42]). The area occupied by a population is most effectively
defined by the ecological boundaries that are important to the population (for example, a particular region
and type of vegetation for a population of beetles, or a particular pond for a population of fish).
The geographic range and distribution of populations (i.e., their spatial structure) represent key factors
in analyzing population diversity because they give an indication of likelihood of movement of organisms
between populations and subsequent genetic and demographic interchange. Similarly, an estimate of the
overall population size provides a measure of the potential genetic diversity within the population; large
populations usually represent larger gene pools and hence greater potential diversity (see Genetic diversity2).
Isolated populations, with very low levels of interchange, show high levels of genetic divergence (Hunter,
2002: 145[42]), and exhibit unique adaptations to the biotic and abiotic characteristics of their habitat. The
genetic diversity of some groups that generally do not disperse well - such as amphibians, mollusks, and
some herbaceous plants - may be mostly restricted to local populations (Avise, 1994[4]). For this reason,
range retractions of species can lead to loss of local populations and the genetic diversity they hold. Loss of
isolated populations along with their unique component of genetic variation is considered by some scientists
to be one of the greatest but most overlooked tragedies of the biodiversity crisis (Ehrlich & Raven 1969[23]).
Populations can be categorized according to the level of divergence between them. Isolated and genetically
distinct populations of a single species may be referred to as subspecies according to some (but not all) species
concepts. Populations that show less genetic divergence might be recognized as variants or races. However,
the distinctions between subspecies and other categories can be somewhat arbitrary (see Species diversity
(Chapter 6)).
A species that is ecologically linked to a specialized, patchy habitat may likely assume the patchy dis-
tribution of the habitat itself, with several different populations distributed at different distances from each
other. This is the case, for example, for species that live in wetlands, alpine zones on mountaintops, par-
ticular soil types or forest types, springs, and many other comparable situations. Individual organisms may
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2"'Genetic Diversity"
41
42 CHAPTER 12. POPULATION DIVERSITY
periodically disperse from one population to another, facilitating genetic exchange between the populations.
This group of different but interlinked populations, with each different population located in its own, discrete
patch of habitat, is called a metapopulation.
There may be quite different levels of dispersal between the constituent populations of a metapopulation.
For example, a large or overcrowded population patch is unlikely to be able to support much immigration
from neighboring populations; it can, however, act as a source of dispersing individuals that will move away
to join other populations or create new ones. In contrast, a small population is unlikely to have a high
degree of emigration; instead, it can receive a high degree of immigration. A population that requires net
immigration in order to sustain itself acts as a sink. The extent of genetic exchange between source and
sink populations depends, therefore, on the size of the populations, the carrying capacity of the habitats
where the populations are found, and the ability of individuals to move between habitats. Consequently,
understanding how the patches and their constituent populations are arranged within the metapopulation,
and the ease with which individuals are able to move among them is key to describing the population diversity
and conserving the species. For more discussion, see the module on Metapopulations.
Chapter 13
Biogeographic Diversityl
Biogeography is "the study of the distribution of organisms in space and through time". Analyses of
the patterns of biogeography can be divided into the two fields of historical biogeography and ecological
biogeography (Wiley, 1981 [109]).
Historical biogeography examines past events in the geological history of the Earth and uses these to
explain patterns in the spatial and temporal distributions of organisms (usually species or higher taxonomic
ranks). For example, an explanation of the distribution of closely related groups of organisms in Africa
and South America is based on the understanding that these two land masses were formerly connected
as part of a single land mass (Gondwana). The ancestors of those related species which are now found
in Africa and South America are assumed to have had a cosmopolitan distribution across both continents
when they were connected. Following the separation of the continents by the process of plate tectonics,
the isolated populations are assumed to have undergone allopatric speciation (i.e., speciation achieved
between populations that are completely geographically separate). This separation resulted in the closely
related groups of species on the now separate continents. Clearly, an understanding of the systematics of
the groups of organisms (i.e., the evolutionary relationships that exists between the species) is an integral
part of these historical biogeographic analyses.
The same historical biogeographic hypotheses can be applied to the spatial and temporal distributions
of marine biota. For example, the biogeography of fishes from different ocean basins has been shown to
be associated with the geological evolution of these ocean basins (see Stiassny and Harrison, 2000[95] for
examples with references). However, we cannot assume that all existing distribution patterns are solely the
product of these past geological processes. It is evident, for example, that the existing marine fauna of the
Mediterranean is a product of the complex geological history of this marine basin, involving separation from
the Indian and Atlantic Oceans, periods of extensive desiccation followed by flooding and recolonization from
the Atlantic (Por, 1989[76]). However, there is also good evidence that the eastern end of the Mediterranean
has been colonized more recently by species that have dispersed from the Red Sea via the Suez canal.
Thus, the field of ecological biogeography first examines the dispersal of organisms (usually individuals
or populations) and the mechanisms that influence this dispersal, and then uses this information to explain
the spatial distribution patterns of these organisms. For further discussion see the module on "Biogeography"
and see Wiley, 1981 [109], and Humphries and Parenti, 1999[39].
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44 CHAPTER 13. BIOGEOGRAPHIC DIVERSITY
Chapter 14
Community Diversity'
A community comprises the populations of different species that naturally occur and interact in a particular
environment. Some communities are relatively small in scale and may have well-defined boundaries. Some
examples are: species found in or around a desert spring, the collection of species associated with ripening
figs in a tropical forest, those clustered around a hydrothermal vent on the ocean floor, those in the spray
zone of a waterfall, or under warm stones in the alpine zone on a mountaintop. Other communities are
larger, more complex, and may be less clearly defined, such as old-growth forests of the northwest coast of
North America, lowland fen communities of the British Isles, or the community of freshwater species of Lake
Baikal.
Sometimes biologists apply the term "community" to a subset of organisms within a larger community.
For example, some biologists may refer to the "community" of species specialized for living and feeding
entirely in the forest canopy, whereas other biologists may refer to this as part of a larger forest community.
This larger forest community includes those species living in the canopy, those on the forest floor, and those
moving between these two habitats, as well as the functional interrelationships between all of these. Similarly,
some biologists working on ecosystem management might distinguish between the community of species that
are endemic to an area (e.g. species that are endemic to an island) as well as those "exotic" species that
have been introduced to that area. The introduced species form part of the larger, modified community of
the area, but might not be considered as part of the regions original and distinctive community.
Communities are frequently classified by their overall appearance, or physiognomy. For example, coral
reef communities are classified according to the appearance of the reefs where they are located, i.e., fringing
reef communities, barrier reef communities, and atoll communities. Similarly, different stream communities
may be classified by the physical characteristics of that part of the stream where the community is located,
such as riffle zone communities and pool communities. However, one of the easiest, and hence most frequent
methods of community classification is based on the dominant types of species present for example, intertidal
mussel bed communities, Ponderosa pine forest communities of the Pacific northwest region of the U.S., or
Mediterranean scrubland communities. Multivariate statistics provide more complex methods for diagnosing
communities, for example, by arranging species on coordinate axes (e.g., x-y axes) that represent gradients in
environmental factors such as temperature or humidity. For more information, see the module on "Natural
communities in space and time."
The factors that determine the diversity of a community are extremely complex. There are many theories
on what these factors are and how they determine community and ecosystem diversity. Environmental
factors, such as temperature, precipitation, sunlight, and the availability of inorganic and organic nutrients
are very important in shaping communities and ecosystems. Hunter (2002: 81)[43] notes that, generally
speaking, organisms can persist and evolve in places where there are sufficient environmental resources for the
organisms to channel energy into growth and reproduction rather than simply the metabolic requirements
for survival. In other words, organisms are less likely to thrive in a harsh environment with low energy
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46 CHAPTER 14. COMMUNITY DIVERSITY
resources. One way of measuring community diversity is to examine the energy flow through food webs that
unite the species within the community; the extent of community diversity can be measured by the number
of links in the food web. However, in practice, it can be very difficult to quantify the functional interactions
between the species within a community. It is easier to measure the genetic diversity of the populations in
the community, and to count the numbers of species present, and use these measures of genetic diversity
and species richness as proxies for describing the functional diversity of the community. The evolutionary
or taxonomic diversity of the species present is another way of measuring the diversity of a community, for
application to conservation biology.
Chapter 15
Ecoregionsi
Since the 1980s, there has been an increasing tendency to map biodiversity over "ecosystem regions" or
"ecoregions". An ecoregion is "a relatively large unit of land or water containing a geographically distinct
assemblage of species, natural communities, and environmental conditions" (WWF, 1999[28]); thus, the
ecosystems within an ecoregion have certain distinct characters in common (Bailey, 1998a[6]). Several
standard methods of classifying ecoregions have been developed, with climate, altitude, and predominant
vegetation being important criteria (Stein et al., 2000 [93]). Bailey's (1983, 1998a, b) classification is one
of the most widely adopted. It is a hierarchical system with four levels: domains, divisions, provinces and
sections.
Domains are the largest geographic levels and are defined by climate, e.g., polar domain, dry domain,
or humid tropical domain. Domains are split into smaller divisions that are defined according climate
and vegetation, and the divisions are split into smaller provinces that are usually defined by their major
plant formations. Some divisions also include varieties of "mountain provinces". These generally have a
similar climatic regime to the neighboring lowlands but show some altitudinal zonation, and they are defined
according to the types of zonation present. Provinces are divided into sections, which are defined by the
landforms present.
Because ecoregions are defined by their shared biotic and abiotic characteristics, they represent practical
units on which to base conservation planning. Moreover, the hierarchical nature of Bailey's ecoregion classifi-
cation allows for conservation management to be planned and implemented at a variety of geographical levels,
from small scale programs focused on discrete sections, to much larger national or international projects that
target divisions. Olson and Dinerstein (2002[72]) identified 238 terrestrial or aquatic ecoregions called the
"Global 200" that they considered to be priorities for global conservation. These ecoregions were selected
because they harbor exceptional biodiversity and are representative of the variety of Earths ecosystems.
For further discussion of ecoregions see the modules on Landscape ecology and Conservation planning on a
regional scale.
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48 CHAPTER 15. ECOREGIONS
Chapter 16
Exteinction
Extinction (the complete disappearance of a species from Earth) is an important part of the evolution of
life on Earth. The current diversity of species is a product of the processes of extinction and speciation
throughout the previous 3.8 billion year history of life. Raup (1991 [78]) assumed that there might be 40
million species alive today, but between 5 and 50 billion species have lived at some time during the history of
the Earth. Therefore, Raup estimated that 99.9% of all the life that has existed on Earth is now extinct); a
species is assumed to be extinct when there is no reasonable doubt that the last individual has died (IUCN,
2002[29]). However, extinction has not occurred at a constant pace through the Earth's history. There have
been at least five periods when there has been a sudden increase in the rate of extinction, such that the rate
has at least doubled, and the extinctions have included representatives from many different taxonomic groups
of plants and animals; these events are called mass extinctions. The timing of these mass extinctions is
shown in Figure 16.1.
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50
50 CHAPTER 16. EXTINCTION
ThaMAJoE1NC'noN EvENr5
Er Period Epoch =Approximate Maor axln ion eventL'
duration of
Era, Period,
or Ep.c~h
(millions of
years futrs
CENOZOIC Qualernary Holocne ,rlt 0.01 arnfjrvl 2~~ii
Tertiary Pliooerke 1 -5U
Oligocenie 24-37
Eoceri 37-58
P______ ____ _ P ere 58--65 rn
MESOZOIC CrtaDOJ ______ 65-144 Omam -
Jurassic ______144-206 MSO r tn ; ~iir6, dc 6(- )a~
__ Tra~s~ _______208-24-5
(Car 286ro ° -32
Devon~ianroa 32600 2 I
Siluriani 408-440n(ad F
Cambir n _____ 605-670
PRECANRIAN 570-450
"Many smaIer etiuidioni events are
not indicated-. see Raup (1991: fig. 4-1 for
Figure 16.1
Each of the first five mass extinctions shown in Figure 16.1 represents a significant loss of biodiversity -
but recovery has been good on a geologic time scale. Mass extinctions are apparently followed by a sudden
burst of evolutionary diversification on the part of the remaining species, presumably because the surviving
species started using habitats and resources that were previously "occupied" by more competitively successful
species that went extinct. However, this does not mean that the recoveries from mass extinction have been
rapid; they have usually required some tens of millions of years (Jablonski, 1995[49]).
It is hypothesized that we are currently on the brink of a "sixth mass extinction," but one that differs
from previous events. The five other mass extinctions predated humans and were probably the ultimate
products of some physical process (e.g. climate change through meteor impacts), rather than the direct
consequence of the action of some other species. In contrast, the sixth mass extinction is the product of
human activity over the last several hundred, or even several thousand years. These mass extinctions, and
their historic and modern consequences are discussed in more detail in the modules on Historical perspectives
on extinction and the current biodiversity crisis, and Ecological consequences of extinctions..
Chapter 17
Landscape Diversity
A landscape is "a mosaic of heterogeneous land forms, vegetation types, and land uses" (Urban et al.,
1987[101]). Therefore, assemblages of different ecosystems (the physical environments and the species that
inhabit them, including humans) create landscapes on Earth. Although there is no standard definition of
the size of a landscape, they are usually in the hundred or thousands of square miles.
Species composition and population viability are often affected by the structure of the landscape; for
example, the size, shape, and connectivity of individual patches of ecosystems within the landscape (Noss,
1990[71]). Conservation management should be directed at whole landscapes to ensure the survival of
species that range widely across different ecosystems (e.g., jaguars, quetzals, species of plants that have
widely dispersed pollen and seeds) (Hunter, 2002: 83-85, 268-270 ([42])).
Diversity within and between landscapes depends on local and regional variations in environmental con-
ditions, as well as the species supported by those environments. Landscape diversity is often incorporated
into descriptions "ecoregions (Chapter 15),"
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52 CHAPTER 17. LANDSCAPE DIVERSITY
Chapter 18
Ecological Value'
Natural communities are finely-tuned systems, where each species has an ecological value to the other
species that are part of that ecosystem. Species diversity increases an ecosystem's stability and resilience, in
particular its ability to adapt and respond to changing environmental conditions. If a certain amount, or type
(such as a keystone species) of species are lost, eventually it leads to the loss of ecosystem function. Many
ecosystems though have built-in redundancies so that two or more species' functions may overlap. Because of
these redundancies, several changes in the number or type of species may not impact an ecosystem. However,
not all species within an ecosystem are of the same importance. Species that are important due to their sheer
numbers are often called dominant species. These species make up the most biomass of an ecosystem.
Species that have important ecological roles that are greater than one would expect based on their abundance
are called keystone species. These species are often central to the structure of an ecosystem, removal of
one or several keystone species may have consequences immediately, or decades or centuries later (Jackson et
al. 2001 [?]). Ecosystems are complex and difficult to study, thus it is often difficult to predict which species
are keystone species. The impact of removing an individual or several keystone species from kelp forests in
the Pacific is examined in Example 18.1 (Northern Pacific Kelp Forests).
Example 18.1: Northern Pacific Kelp Forests
Kelp forests, as their name suggests, are dominated by kelp, a brown seaweed of the family
Laminariales. They are found in shallow, rocky habitats from temperate to subarctic regions, and
are important ecosystems for many commercially valuable fish and invertebrates.
Vast forests of kelp and other marine plants existed in the northern Pacific Ocean prior to the
18th century. The kelp was eaten by herbivores such as sea urchins (Family Strongylocentrotidae),
which in turn were preyed upon by predators such as sea otters (Enhydra lutris). Hunting during
the 18th and 19th centuries brought sea otters to the brink of extinction. In the absence of sea
otters, sea urchin populations burgeoned and grazed down the kelp forests, at the extreme creating
"urchin barrens," where the kelp was completely eradicated. Other species dependent on kelp
(such as red abalone Haliotis rufescens) were affected too. Legal protection of sea otters in the 20th
century led to partial recovery of the system.
More recently sea otter populations in Alaska seem to be threatened by increased predation
from killer whales (Orcinus orca) (Estes et al. 1998[?]). It appears that whales may have shifted
their diet to sea otters when populations of their preferred prey, Stellar sea lions (Arctocephalus
townsendi) and Harbor seals (Phoca vitulina) declined. The exact reason for the decline in the
sea lion and seal populations is still unclear, but appears to be due to declines in their prey in
combination with increased fishing and higher ocean temperatures. As a result of the loss of sea
otters, increased sea urchin populations are grazing down kelp beds again.
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54 CHAPTER 18. ECOLOGICAL VALUE
Example 18.2: Southern Californian Kelp Forests
Interestingly, a similar scenario in kelp forests in Southern California did not show immediate
effects after the disappearance of sea otters from the ecosystem. This is because the system was
more diverse initially. Other predators (California sheephead fish, Semicossyphus pulcher, and spiny
lobsters, Panulirus interruptus) and competitors (abalone Haliotis spp) of the sea urchin helped
maintain the system. However, when these predators and competitors were over-harvested as well
in the 1950s, the kelp forests declined drastically as sea urchin populations boomed.
In the 1970s and 1980s, a sea urchin fishery developed which then enabled the kelp forest to
recover. However, it left a system with little diversity. The interrelationships among these species
and the changes that reverberate through systems as species are removed are mirrored in other
ecosystems on the planet, both aquatic and terrestrial.
As this example illustrates, biodiversity is incredibly complex and conservation efforts cannot
focus on just one species or even on events of the recent past.
GLOSSARY
55
Glossary
A albedo
the amount of solar radiation reflected by
a surface
allopatric speciation
speciation achieved between populations
that are completely geographically
separated (their ranges do not overlap or
are not contiguous).
Alpha diversity
the diversity within a particular area or
ecosystem; usually expressed by the
number of species (i.e., species richness)
in that ecosystem
Area of endemism
an areas which has a high proportion of
endemic species (i.e., species with
distributions that are naturally restricted
to that region)
B bequest value
the value of knowing something will be
there for future generations
Beta diversity
a comparison of of diversity between
ecosystems, usually measured as the
amount of species change between the
ecosystems
Biodiversity coldspots
areas that have relatively low biological
diversity but are also experiencing a high
rate of habitat loss
Biodiversity hotspots
in general terms these are areas that have
high levels of endemism (and hence
diversity) but which are also experiencing
a high rate of loss of habitat. This
concept was originally developed for
terrestrial ecosystems. A terrestrial
biodiversity hotspot is an area that has
at least 0.5%, or 1,500 of the worlds ca.
300,000 species of green plants
(Viridiplantae), and that has lost at least
70% of its primary vegetation (Myers et
al., 2000[?]). Marine biodiversity
hotspots have been defined for coral reefs,
based on measurements of relative
endemism of multiple taxa (species of
corals, snails, lobsters, fishes) within a
region and the relative level of threat to
that region (Roberts et al., 2002[?])
Biodiversity
the variety of life on Earth at all its
levels, from genes to ecosystems, and the
ecological and evolutionary processes that
sustain it
biogeography
the study of the distribution of organisms
in space and through time
Biological species concept
a species is a group of interbreeding
natural populations unable to
successfully mate or reproduce with other
such groups, and which occupies a
specific niche in nature (Mayr, 1982;
Bisby and Coddington, 1995).
C Community
the populations of different species that
naturally occur and interact in a
particular environment
Community
the populations of different species that
naturally occur and interact in a
particular environment.
D Demography
the statistical characteristics of the
population such as size, density, birth
and death rates, distribution, and
movement or migration.
direct use value
56
GLOSSARY
refers to products or goods which are
consumed directly such as food or timber
dominant species
species that are important due to their
sheer numbers in an ecosystem
E Ecological biogeography:
the study of the dispersal of organisms
(usually individuals or populations) and
the mechanisms that influence this
dispersal, and the use of this information
to explain spatial distribution patterns
ecological value
the values that each species has as part of
an ecosystem
Ecoregion
a relatively large unit of land or water
containing a geographically distinct
assemblage of species, natural
communites, and environmental
conditions (WWF, 1999[?])
Ecoregions
a relatively large unit of land or water
containing a geographically distinct
assemblage of species, natural
communities, and environmental
conditions (WWF, 1999[?]). The
ecosystems within an ecoregion have
certain distinct characters in common
(Bailey, 1998a[?]).
Ecosystem
a community plus the physical
environment that it occupies at a given
time
ecosystem
a community plus the physical
environment that it occupies at a given
time.
Ecosystem
Endemic species
those species whose distributions are
naturally restricted to a defined region
evapotranspiration
is the process whereby water is absorbed
from soil by vegetation and then released
back into the atmosphere
Evolutionary significant unit
a group of organisms that has undergone
significant genetic divergence from other
groups of the same species. Identification
of ESUs is based on natural history
information, range and distribution data,
and results from analyses of
morphometrics, cytogenetics, allozymes
and nuclear and mitochondrial DNA.
Concordance of those data, and the
indication of significant genetic distance
between sympatric groups of organisms,
are critical for establishing an ESU.
existence value
the value of knowing something exists even
if you will never use it or see it
Extinct
a species is assumed to be extinct when
there is no reasonable doubt that the last
individual has died (IUCN, 2002[?])
Extinction
the complete disappearance of a species
from Earth
G Gamma diversity
a measure of the overall diversity within a
large region. Geographic-scale species
diversity according to Hunter (2002: 448)
([42])
Genetic Diversity
refers to any variation in the nucleotides,
genes, chromosomes, or whole genomes of
organisms.
H Historical biogeography
the study of events in the geological
history of the Earth and their use to
explain patterns in the spatial and
temporal distributions of organisms
(usually species or higher taxonomic
ranks)
I indirect use value
refers to the services that support the
products that are consumed, this includes
ecosystems functions like nutrient cycling
K keystone species
GLOSSARY
57
species that have important ecological
roles that are greater than one would
expect based on their abundance
L Landscape
a mosaic of heterogeneous land forms,
vegetation types, and land uses (Urban et
al., 1987[?])
Landscapes
a mosaic of heterogeneous land forms,
vegetation types, and land uses (Urban et
al., 1987[?]).
M Marine Biodiversity hotspots
Mass extinction
a period when there is a sudden increase
in the rate of extinction, such that the
rate at least doubles, and the extinctions
include representatives from many
different taxonomic groups of plants and
animals
Metapopulation
a group of different but interlinked
populations, with each different
population located in its own, discrete
patch of habitat
Morphological species concept
species are the smallest natural
populations permanently separated from
each other by a distinct discontinuity in
the series of biotype (Du Rietz, 1930;
Bisby and Coddington, 1995).
N non use or passive value
refers to the value for things that we don't
use but would feel a loss if they were to
disappear
O Orobiome
a mountainous environment or landscape
with its constituent ecosystems
Orogenesis
the process of mountain building.
occupying contiguous but not overlapping
ranges.
Photosynthesis
the formation of carbohydrates from
carbon dioxide and water, through the
action of light energy on a light-sensitive
pigment, such as chlorophyll, and usually
resulting in the production of oxygen
Phylogenetic diversity
the evolutionary relatedness of the species
present in an area.
Phylogenetic species concept
a species is the smallest group of
organisms that is diagnosably [that is,
identifiably] distinct from other such
clusters and within which there is a
parental pattern of ancestry and descent
(Cracraft, 1983; Bisby and Coddington,
1995).
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".
Population
1. a group of individuals of the same species
that share aspects of their demography or
genetics more closely with each other
than with other groups of individuals of
that species
2. A population may also be defined as a
group of individuals of the same species
occupying a defined area at the same
time (Hunter, 2002: 144[?])
potential or option value
refers to the use that something may have
in the future
S Sink
a population patch, in a metpopulation
that does not have a high degree of
emigration outside its boundaries but,
instead, requires net immigration in order
to sustain itself
Source
P Parapatric
58
GLOSSARY
a population patch, in a metapopulation,
from which individuals disperse to other
population patches or create new ones
Species diversity
the number of different species in a
particular area (i.e., species richness)
weighted by some measure of abundance
such as number of individuals or biomass.
Species evenness
the relative abundance with which each
species are represented in an area.
Species richness
the number of different species in a
particular area
Species richness
the number of different species in a
particular area.
surface roughness
the average vertical relief and small-scale
irregularities of a surface
Sympatric
occupying the same geographic area.
T Terrestrial Biodiversity hotspots
v watersheds
land areas drained by a river and its
tributaries
wetlands
areas where water is present at or near the
surface of the soil or within the root zone,
all year or for a period of time during the
year, and where the vegetation is adapted
to these conditions
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66
INDEX
Index of Keywords and Terms
Keywords are listed by the section with that keyword (page numbers are in parentheses). Keywords
do not necessarily appear in the text of the page. They are merely associated with that section. Ex.
apples, j 1.1 (1) Terms are referenced by the page they appear on. Ex. apples, 1
A albedo, 2
allopatric speciation, 43
Alpha diversity, 31
B bequest value, 33
beta diversity, 31
biodiversity, @ 2(5), 5, 5, @ 3(7), @ 4(9), @ 5(11),
@ 6(25), @ 7(31), @ 10(37), @ 11(39), @ 12(41),
@ 13(43), @ 14(45), @ 15(47), @ 16(49), @ 17(51)
biodiversity coldspots, 8
biodiversity hotspots, 7
Biogeography, 43
biological species concept, 26
C collector's curve, 15
communities, 9
community, 45
D demography, 41
direct use value, 33
dominant species, 53
E E, 25
ecological biogeography, 43
ecological value, 53
ecoregion, 47
ecoregions, 9
ecosystem, 25, 39
ecosystems, 7, 9, 31, 45
endemic species, 7
evapotranspiration, 2, 2
evolutionary significant unit, 26
existence value, 33
extinct, 49
Extinction, 49
G Gamma diversity, 31
genetic diversity, 9
H H, 25
Historical biogeography, 43
indirect use value, 33
K keystone species, 53
L landscape, 51
landscapes, 9
M Marine biodiversity hotspots, 8
mass extinctions, 49
metapopulation, 42
morphological species concept, 26, 26
N non-use or passive values, 33
O orobiomes, 7
orogenesis, 5
P parapatric, 26
Photosynthesis, 37
phylogenetic diversity, 25
phylogenetic species concept, 26
physiognomy, 45
plate tectonics, 5
population, 41, 41
population diversity, 9
Potential or Option value, 33
R races., 41
radiation balance, 2
S sink, 42
source, 42
species diversity, 9, 25
species evenness, 25
species richness, 7, 25, 31
surface roughness, 2, 2
sympatric, 26
T terrestrial biodiversity hotspot, 7
V variants, 41
W watersheds, 2
wetlands, 3
I incipient species, 26
ATTRIBUTIONS
67
Attributions
Collection: What is Biodiversity
Edited by: Nora Bynum
URL: http://cnx.org/content/col10639/1.1/
License: http://creativecommons.org/licenses/by/3.0/
Module: "Global Processes"
By: Melina Laverty, Eleanor Sterling
URL: http://cnx.org/content/m12159/1.1/
Pages: 1-4
Copyright: Melina Laverty, Eleanor Sterling
License: http://creativecommons.org/licenses/by/1.0
Module: "Definition of Biodiversity"
By: Ian Harrison, Melina Laverty, Eleanor Sterling
URL: http://cnx.org/content/m12151/1.2/
Pages: 5-6
Copyright: Ian Harrison, Melina Laverty, Eleanor Sterling
License: http://creativecommons.org/licenses/by/1.0
Module: "Spatial Gradients in Biodiversity"
By: Ian Harrison, Melina Laverty, Eleanor Sterling
URL: http://cnx.org/content/m12173/1.2/
Pages: 7-8
Copyright: Ian Harrison, Melina Laverty, Eleanor Sterling
License: http://creativecommons.org/licenses/by/1.0
Module: "Introduction to the Biodiversity Hierarchy"
By: Ian Harrison, Melina Laverty, Eleanor Sterling
URL: http://cnx.org/content/m12162/1.2/
Page: 9
Copyright: Ian Harrison, Melina Laverty, Eleanor Sterling
License: http://creativecommons.org/licenses/by/1.0
Module: "What is Biodiversity? A comparison of spider communities"
By: James Gibbs
URL: http://cnx.org/content/m12179/1.1/
Pages: 11-24
Copyright: James Gibbs
License: http://creativecommons.org/licenses/by/1.0
Module: "Species Diversity"
By: Ian Harrison, Melina Laverty, Eleanor Sterling
URL: http://cnx.org/content/m12174/1.3/
Pages: 25-29
Copyright: Jan Harrison, Melina Laverty, Eleanor Sterling
License: http://creativecommons.org/licenses/by/1.0
68
ATTRIBUTIONS
Module: "Alpha, Beta, and Gamma Diversity"
By: Ian Harrison, Melina Laverty, Eleanor Sterling
URL: http://cnx.org/content/m12147/1.2/
Pages: 31-32
Copyright: Ian Harrison, Melina Laverty, Eleanor Sterling
License: http://creativecommons.org/licenses/by/1.0
Module: "Introduction to Utilitarian Valuation of Biodiversity"
By: Melina Laverty, Eleanor Sterling
URL: http://cnx.org/content/m12164/1.2/
Pages: 33-34
Copyright: Melina Laverty, Eleanor Sterling
License: http://creativecommons.org/licenses/by/1.0
Module: "Biodiversity over Time"
By: Robert Ahlfinger
URL: http://cnx.org/content/m12148/1.2/
Page: 35
Copyright: Robert Ahlfinger
License: http://creativecommons.org/licenses/by/1.0
Module: "A Brief History of Life on Earth"
By: Ian Harrison, Melina Laverty, Eleanor Sterling
URL: http://cnx.org/content/m12146/1.2/
Pages: 37-38
Copyright: Ian Harrison, Melina Laverty, Eleanor Sterling
License: http://creativecommons.org/licenses/by/1.0
Module: "Ecosystem Diversity"
By: Ian Harrison, Melina Laverty, Eleanor Sterling
URL: http://cnx.org/content/m12156/1.2/
Pages: 39-40
Copyright: Ian Harrison, Melina Laverty, Eleanor Sterling
License: http://creativecommons.org/licenses/by/1.0
Module: "Population Diversity"
By: Ian Harrison, Melina Laverty, Eleanor Sterling
URL: http://cnx.org/content/m12171/1.2/
Pages: 41-42
Copyright: Ian Harrison, Melina Laverty, Eleanor Sterling
License: http://creativecommons.org/licenses/by/1.0
Module: "Biogeographic Diversity"
By: Ian Harrison, Melina Laverty, Eleanor Sterling
URL: http://cnx.org/content/m12149/1.2/
Page: 43
Copyright: Ian Harrison, Melina Laverty, Eleanor Sterling
License: http://creativecommons.org/licenses/by/1.0
Module: " Community Diversity"
By: Jan Harrison, Melina Laverty, Eleanor Sterling
URL: http://enx.org/content/m12150/1.2/
Pages: 45-46
Copyright: Jan Harrison, Melina Laverty, Eleanor Sterling
License: http://creativecommons.org/licenses/by/1.0
ATTRIBUTIONS
69
Module: "Ecoregions"
By: Ian Harrison, Melina Laverty, Eleanor Sterling
URL: http://cnx.org/content/m12155/1.2/
Page: 47
Copyright: Ian Harrison, Melina Laverty, Eleanor Sterling
License: http://creativecommons.org/licenses/by/1.0
Module: "Extinction"
By: Ian Harrison, Melina Laverty, Eleanor Sterling
URL: http://cnx.org/content/m12157/1.2/
Pages: 49-50
Copyright: Ian Harrison, Melina Laverty, Eleanor Sterling
License: http://creativecommons.org/licenses/by/1.0
Module: "Landscape Diversity"
By: Ian Harrison, Melina Laverty, Eleanor Sterling
URL: http://cnx.org/content/m12165/1.2/
Page: 51
Copyright: Ian Harrison, Melina Laverty, Eleanor Sterling
License: http://creativecommons.org/licenses/by/1.0
Module: "Ecological Value"
By: Melina Laverty, Eleanor Sterling
URL: http://cnx.org/content/m12154/1.2/
Pages: 53-54
Copyright: Melina Laverty, Eleanor Sterling
License: http://creativecommons.org/licenses/by/1.0
What is Biodiversity
This collection provides an overview of what is meant by the term 'biodiversity,' and how we measure it.
The collection reviews the different levels of biodiversity, or the 'biodiversity hierarchy' including: genetic
and phenotypic diversity; population diversity; species diversity; community diversity; ecosystem diversity;
landscape diversity; and historical and ecological biogeographic diversity. Brief definitions of populations,
species, communities, and ecosystems are provided, with some introductory discussion of different types
of 'species concepts.' The collection defines the terms 'species richness' and 'species evenness' as methods
for measuring species diversity, and it discusses the use of species richness as a surrogate for describing
overall global biodiversity. The collection reviews the distribution of biodiversity in space, explaining the
definitions of alpha, beta and gamma diversity for measuring diversity within and between ecosystems. The
environmental factors that affect these patterns of spatial diversity are briefly discussed. The collection also
includes a brief review of the different ways by which assessments of spatial diversity are used for conservation
planning and management (e.g., based on ecoregions, or biodiversity hotspots and coldspots). The collection
concludes with a brief discussion of diversity over geological time.
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