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Andrew John Hamilton

University of Melbourne

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 Journal of Environmental Management 75 (2005) 89–92
www.elsevier.com/locate/jenvman

keragaman spesies atau keanekaragaman hayati?

Andrew J. Hamilton *
Industri Primer Penelitian Victoria-Knox lapangan, Private Bag 15, Ferntree Gully Pengiriman Pusat, Victoria 3156, Australia

Menerima 8 Maret 2004; direvisi Oktober 2004 27; diterima November 2004 4

Abstrak

keanekaragaman spesies dan keanekaragaman hayati secara luas digunakan istilah dalam ekologi dan pengelolaan sumber daya alam. Meskipun demikian, mereka tidak mudah didefinisikan dan penulis yang

berbeda berlaku istilah-istilah ini dengan berbagai konotasi. Keanekaragaman hayati Istilah, khususnya, memiliki kehormatan yang meragukan yang banyak digunakan tapi jarang didefinisikan. Apakah ini hanya jumlah

spesies atau itu sesuatu yang lebih? Di sini saya mempertimbangkan apa istilah-istilah ini mungkin benar-benar berarti dan nilai mereka. Saya juga secara singkat y membahas alasan untuk mempelajari dan melindungi

keragaman spesies atau keanekaragaman hayati. Crown Copyright q 2005 Diterbitkan oleh Elsevier Ltd All rights reserved.

Kata kunci: keanekaragaman hayati; keragaman ekologi; keanekaragaman spesies; kekayaan spesies

1. Perkenalan
When considering the basic structure of biological systems such as
communities or ecosystems, two fundamental 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 scientific 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 define 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 difficult to define and measure,
and this may in fact reflect 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

* Tel.: C 61 3 9210 9282; fax: C 61 3 9800 3521. theory, such as grammatical diversity in manuscripts (e.g. Margalef, 1958;
E-mail address: andrew.hamilton@dpi.vic.gov.au 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
90 A.J. Hamilton / Journal of Environmental Management 75 (2005) 89–92
and Weaver, 1949 ). The similarities between artificial 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 equitability (the distribution of
abundance among the species). Different diversity indices apportion
different relative weights to these properties, and the definition of evenness
itself varies. Justifications 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 ð N i= N Þ 2, where N i is the number of
individuals in the i th species).
Perhaps the most meaningful but seldom used index, not a diversity
index sensu stricto, is PIE (probability of interspecific encounter), which
attempts to calculate, for a hypothetical individual, the proportion of
encounters that will be with another species ( Hurlbert, 1971 ).
" #
2
Ni
PIE Z N
NK11KX N many propound a much broader definition such as the ‘full variety of life on
i
where N and N i are as defined 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 randommethod of
wind dispersal of pollen ( Hurlbert, 1971 ). It should be noted that Simpson’s
(1949) index is closely related to PIE.
Describing species diversity as a single value compromises 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 MacArthur’s 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) defined 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 diversity 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 definitions of biodiversity and most are vague,
which probably reflects 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
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 definitions of biodiversity. The International Convention on
Biological Diversity (2003)
uses the following definition:
“Biological diversity” [biodiversity] means the variability among living
organisms from all sources including,
inter alia, terrestrial, marine and other aquatic ecosystems 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 word’s 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 terminology
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) 89–92
( 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 justification for disciplines such as systematics, which
traditionally had difficulty convincing funding bodies of the value of their
work. This somewhat disingenuous origin in the scientific 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 definition 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-ecosystems. 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 Benefits
Index (BBI) is an extension/modification of the habitat hectares index ( Oliver
and Parkes, 2003 ). Neither of these metrics have strict ecological meanings
( c.f. Hurlbert’s 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
significance of the mathematical manipulations applied to components,
summation for habitat hectares and multiplication for BBI, used to obtain
the metric are not explained or justified.
4. Justifications 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 defined 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 fulfil very similar functions, and thus loss of some
of these would have little
91
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
reckoning 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 problem—how do we
define ecosystem functioning? One definition 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 justifications for
protecting biodiversity. There is a danger that the public or industry will
have a suspicious opinion of biodiversity if its benefits are mostly expressed
in terms of reimbursements to humans. Such (perceived) benefits 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 definition and measure being used. Biodiversity,
on the other hand, is an unclear concept, and its value to the ecologist is
questionable. 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.
92 AJ Hamilton / Jurnal Manajemen Lingkungan 75 (2005) 89-92
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