Species diversity or biodiversity?

Andrew J. Hamilton
*
Primary Industries Research VictoriaKnoxeld, Private Bag 15, Ferntree Gully Delivery Centre, Victoria 3156, Australia
Received 8 March 2004; revised 27 October 2004; accepted 4 November 2004
Abstract
Species diversity and biodiversity are widely used terms in ecology and natural resource management. Despite this, they are not easily
dened and different authors apply these terms with varying connotations. The term biodiversity, in particular, has the dubious honour of
being widely used but rarely dened. Is it simply the number of species or is it something more? Here I consider what these terms might really
mean and their value. I also briey discuss the rationale for studying and protecting species diversity or biodiversity.
Crown Copyright q 2005 Published by Elsevier Ltd. All rights reserved.
Keywords: Biodiversity; Ecological diversity; Species diversity; Species richness
1. Introduction
When considering the basic structure of biological
systems such as communities or ecosystems, two funda-
mental parameters are the number of species and the number
of individuals within each of these species. Ecologists have
studied the inter-relationships between these in considerable
depth over many decades (e.g. Fisher 1943; MacArthur,
1957; Hurlbert, 1971; May, 1975; Sugihara, 1980), and in
doing so have primarily been concerned with what they call
species diversity. More recently, a concept known as
biodiversity has been rapidly gaining prominence in
political, management, public, and scientic arenas
(DeLong, 1996, Williams in press). There has been
considerable confusion, however, as to what exactly is
meant by biodiversity, and its connection to more traditional
concepts such as species diversity is unclear. In this paper I
discuss both concepts, identify possible commonalities and
differences, and consider their usefulness.
2. Species diversity: what is it and how do we measure it?
Traditionally, ecologists have been concerned with the
concept of ecological diversity. Species diversity is the most
commonly used representation of ecological diversity, but it
is not the only measure. Niche width and habitat diversity
are also key components of ecological diversity. Niche
width describes the availability of resources to an organism
(or taxon) over spatial and temporal scales. It is the breadth
or diversity of resources used by an individual/taxon
(Magurran, 1988). Habitat diversity measures the structural
complexity of the environment (Mumby, 2001). I will
primarily focus on species diversity, but it should be noted
that many of the methods used to dene species diversity are
also applicable to a degree to niche width and habitat
diversity. The concepts of niche width and habitat diversity
are of utmost importance for understanding ecosystem
function.
Ecologists have found species diversity difcult to dene
and measure, and this may in fact reect the possibility that
it is a non-concept (Hurlbert, 1971). In general, there have
been two approaches to measuring species diversity, both of
which incorporate information on the number of species
(species richness) and the relative abundances of individuals
within each species (species abundance). One method has
been to construct mathematical indices broadly known as
diversity indices; the other involves comparing observed
patterns of species abundance to theoretical species
abundance models.
Many commonly used diversity indices are derived from
information theory, such as grammatical diversity in
manuscripts (e.g. Margalef, 1958; Shannon, 1948; Shannon
0301-4797/$ - see front matter Crown Copyright q 2005 Published by Elsevier Ltd. All rights reserved.
doi:10.1016/j.jenvman.2004.11.012
Journal of Environmental Management 75 (2005) 8992
www.elsevier.com/locate/jenvman
* Tel.: C61 3 9210 9282; fax: C61 3 9800 3521.
E-mail address: andrew.hamilton@dpi.vic.gov.au
and Weaver, 1949). The similarities between articial
information sets, such as letters or words on a page, and
natural communities have not been determined, rather only
assumed or at best, implied (Hurlbert, 1971). Species
diversity indices take two aspects of a community into
account, namely species richness and evenness or equit-
ability (the distribution of abundance among the species).
Different diversity indices apportion different relative
weights to these properties, and the denition of evenness
itself varies. Justications for these weightings are largely
subjective, and consequently diversity indices do not have
clear ecological meaning. Such limitations can be seen
mathematically as well. May (1975) pointed out that
diversity indices are simply moments of the full S(N)
distribution, where S is the number of species and N is the
total number of individuals. Considering this, he suggested
that the variance of the distribution would be a sensible
measure of diversity. He also propounds that if a diversity
index is to be used, then a variance-based index is preferable
(i.e. one that includes
P
N
i
=N
2
, where N
i
is the number of
individuals in the ith species).
Perhaps the most meaningful but seldom used index, not
a diversity index sensu stricto, is PIE (probability of
interspecic encounter), which attempts to calculate, for a
hypothetical individual, the proportion of encounters that
will be with another species (Hurlbert, 1971).
PIE Z
N
N K1

1 K
X
i
N
i
N

2
" #
where N and N
i
are as dened above. PIE assumes that each
individual within a community can encounter or interact
with every other individual, an assumption that would not
hold true in reality. Nonetheless, this simple index has a
clear ecological interpretation. In high PIE communities,
there would be less reliance by species on random processes
in searching behaviour. For example, it is uncommon for
plant species in high PIE communities such as tropical
rainforests to breed by the random method of wind dispersal
of pollen (Hurlbert, 1971). It should be noted that Simpsons
(1949) index is closely related to PIE.
Describing species diversity as a single value compro-
mises much of the detailed structure of a community.
Theoretical rank-abundance models provide a more
complete picture of community structure (May, 1976).
These include, among others, the log normal distribution
(Sugihara, 1980), the geometric series (Motomura, 1932;
May, 1975), the logarithmic series (Fisher et al., 1943), and
MacArthurs broken stick model (MacArthur, 1957; Webb,
1974; May, 1975). The geometric series and the broken
stick describe the respective opposing extremes where a
community is dominated by a few species and where species
are equally abundant. The log series and the log normal
describe intermediate situations, with the former being
closer to geometric and latter to the broken stick. A
pragmatic limitation of such models is that they are not
always amenable to comparisons between communities: the
one model may not adequately describe all communities.
Species diversity, or other forms of diversity for that
matter, can be partitioned across spatial scales. Whittaker
(1960, 1977) dened a hierarchical system whereby point
diversity is the diversity within a microhabitat, a diversity is
that within a homogenous habitat, g diversity is that within
landscape level units such as islands, and 3 diversity
describes diversity at the scale of biogeographical regions.
This multilevel diversity is also called inventory diversity.
Overlaying the concepts of diversity, species or habitat,
described above is the idea of differentiation diversity. This
describes the degree of change in diversity over space, such
as along a transect or between habitats. Whittaker (1960,
1977) outlined three spatial-levels of differentiation diver-
sity that correspond to his inventory diversity: pattern
diversity, b diversity, and d diversity. Of these, b diversity is
the most commonly measured (Mumby, 2001; Vellend,
2001; Crist et al., 2003), and it describes change in diversity
along a transect or the difference between habitats.
3. Biodiversity: what is it and how do we measure it?
There are currently many denitions of biodiversity and
most are vague, which probably reects the uncertainty of
the concept. Some consider it to be synonymous with
species richness (Marc and Canard, 1997; Heywood, 1998),
others see it as species diversity (Bond and Chase, 2002),
whereas many propound a much broader denition such as
the full variety of life on Earth (Takacs, 1996). NRE
(1997) distinguish between native and introduced species,
and others have put extra emphasis on threatened species
(Brockerhoff et al., 2001). DeLong (1996) reviewed and
assessed a large number of denitions of biodiversity. The
International Convention on Biological Diversity (2003)
uses the following denition:
Biological diversity [biodiversity] means the varia-
bility among living organisms from all sources including,
inter alia, terrestrial, marine and other aquatic ecosys-
tems and the ecological complexes of which they are
part; this includes diversity within species, between
species, and of ecosystems.
One way to gain an appreciation of the words most
widely accepted meaning is to consider its evolution. In
1980 the term biological diversity was used to describe what
was presumably species richness (Lovejoy 1980). In the
same year, Norse and McManus (1980) used this terminol-
ogy to describe a concept that incorporated both ecological
diversity and genetic diversity. In 1981 the US Strategy
Conference on Biological Diversity was held. In 1985 the
National Forum on BioDiversity took place, and the use
of the contracted form of the term in the proceedings of
this conference probably initiated its widespread use
A.J. Hamilton / Journal of Environmental Management 75 (2005) 8992 90
(Harper and Hawksworth, 1994). Ghilarov (1996) noted that
the word biodiversity was originally used in political debate
rather than science. But scientists soon adopted it to secure
research funding. In particular, it seemed to provide a
justication for disciplines such as systematics, which
traditionally had difculty convincing funding bodies of the
value of their work. This somewhat disingenuous origin in
the scientic literature is probably responsible for the
current ambiguity of the word, especially its synonymous
treatment with ecological diversity, species diversity, and
species richness.
There are many ways of measuring biodiversity.
Clearly, the indices and models described earlier are
applicable if we consider biodiversity to be synonymous
with species diversity. However, metrics that accommodate
a broader denition of biodiversity have also been
developed (summarised in Williams 2004). For example,
habitat hectares is an index that combines vegetation area
with its quality (Parkes et al., 2003). The index is
primarily designed for the management of agro-ecosys-
tems. Quality parameters include canopy cover, the number
of large trees, number of logs, patch size and proximity to a
stand of native vegetation. The Biodiversity Benets Index
(BBI) is an extension/modication of the habitat hectares
index (Oliver and Parkes, 2003). Neither of these metrics
have strict ecological meanings (c.f. Hurlberts PIE). The
weights assigned to the various components are purely
subjective in the sense that they do not represent actual
ecological processes, and the statistical distributions
associated with each component are not considered.
Moreover, as noted by Williams (2004), the ecological
signicance of the mathematical manipulations applied to
components, summation for habitat hectares and multipli-
cation for BBI, used to obtain the metric are not explained
or justied.
4. Justications for protecting species diversity
or biodiversity
The study of species diversity, or at least species
richness, gives ecologists insights into the stability of
communities (Walker, 1988). The relationship between
species diversity/richness and community stability is quite
complex. Stability can be dened as the ability of a system
to recover to an equilibrium state after disturbance, or
simply persistence of the system (May, 1976). The
diversity-stability hypothesis asserts that species vary in
their traits, and that in a highly diverse (species rich)
system there will be some species that can compensate for
the loss of others after disturbance (Elton, 1958; Pimm,
1984). Thus, species rich systems are more likely to be
considered stable. The species (functional) redundancy
hypothesis, on the other hand, propounds that in most
ecosystems there are several species that full very similar
functions, and thus loss of some of these would have little
impact on the system as a whole (Walker, 1988). There is
empirical evidence supporting both the diversity-stability
(Tilman and Downing, 1994) and species redundancy
(Bellwood et al., 2003) hypotheses. Mathematical reckon-
ing has demonstrated that the stability of any system
increases with the complexity (i.e. number of components)
only until a critical point is reached, at which the system
becomes unstable (Gardner and Ashby, 1970; May, 1972).
Species diversity has often been discussed in relation to
ecosystem functioning. But here we run into a further
problemhow do we dene ecosystem functioning? One
denition is simply the total production of organic matter or
consumption of carbon dioxide, whereas another includes
the synthesis of all compounds that the organisms of a
community contain (Ghilarov, 2000). Furthermore, the
relationship between species diversity and ecosystem
functioning appears to change over different spatial scales
(Bond and Chase, 2002; Jonathan, 2002).
Ghilarov (2000) has argued that more emphasis should
be put on the intrinsic worth of biodiversity as a rationale for
valuing it. He states that if biodiversity has intrinsic value it
could in principle be useless for human needs or for
ecosystem functioning. This is an important argument, as
it removes the anthropocentric theme that underlies many
justications for protecting biodiversity. There is a danger
that the public or industry will have a suspicious opinion of
biodiversity if its benets are mostly expressed in terms of
reimbursements to humans. Such (perceived) benets are
likely to be indirect or even non-existent, and this could
lead to a devaluing or mistrust of biodiversity. People need
to be aware of issues such as the uniqueness of species, the
right of species to exist, and the irreversible nature of
extinction.
5. Conclusion
Species diversity can be a useful tool for the ecologist,
but one must be clear as to the particular denition and
measure being used. Biodiversity, on the other hand, is an
unclear concept, and its value to the ecologist is question-
able. Nonetheless, it is well entrenched in the world of
environmental management and policy, and it may be that it
is a useful tool from a sociological and political perspective,
even if it does have substantial theoretical limitations.
Ultimately, if biodiversity plays a key role in minimising
anthropogenic impacts on nature, then it is indeed valuable
as a word and concept.
Acknowledgements
This paper is dedicated to the late Emily Hamilton, who
provided me with the stimulus to complete it.
A.J. Hamilton / Journal of Environmental Management 75 (2005) 8992 91
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