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Andrew John Hamilton

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Journal of Environmental Management 75 (2005) 89–92

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


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 commonly used representation of ecological diversity, but it is not the only
measure. Niche width and habitat diversity are also key components of
When considering the basic structure of biological systems such as ecological diversity. Niche width describes the availability of resources to
communities or ecosystems, two fundamental parameters are the number an organism (or taxon) over spatial and temporal scales. It is the breadth or
of species and the number of individuals within each of these species. diversity of resources used by an individual/taxon ( Magurran, 1988 ).
Ecologists have studied the inter-relationships between these in Habitat diversity measures the structural complexity of the environment ( Mumby,
considerable depth over many decades (e.g. Fisher 1943; MacArthur, 2001 ). I will primarily focus on species diversity, but it should be noted that
1957; Hurlbert, 1971; May, 1975; Sugihara, 1980 ), and in doing so have many of the methods used to define species diversity are also applicable to
primarily been concerned with what they call species diversity. More a degree to niche width and habitat diversity. The concepts of niche width
recently, a concept known as biodiversity has been rapidly gaining and habitat diversity are of utmost importance for understanding ecosystem
prominence in political, management, public, and scientific arenas ( DeLong, function.
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, Ecologists have found species diversity difficult to define and measure,
and consider their usefulness. 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
2. Species diversity: what is it and how do we measure it? comparing observed patterns of species abundance to theoretical species
abundance models.
Traditionally, ecologists have been concerned with the concept of
ecological diversity. Species diversity is the most

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: Shannon, 1948; Shannon

0301-4797/$ - see front matter Crown Copyright q 2005 Published by Elsevier Ltd. All rights reserved.
90 A.J. Hamilton / Journal of Environmental Management 75 (2005) 89–92

and Weaver, 1949 ). The similarities between artificial information sets, such always amenable to comparisons between communities: the one model
as letters or words on a page, and natural communities have not been may not adequately describe all communities.
determined, rather only assumed or at best, implied ( Hurlbert, 1971 ). Species diversity, or other forms of diversity for that matter, can be
Species diversity indices take two aspects of a community into account, partitioned across spatial scales. Whittaker (1960, 1977) defined a
namely species richness and evenness or equitability (the distribution of hierarchical system whereby point diversity is the diversity within a
abundance among the species). Different diversity indices apportion microhabitat, a diversity is that within a homogenous habitat, g diversity is
different relative weights to these properties, and the definition of evenness that within landscape level units such as islands, and 3 diversity describes
itself varies. Justifications for these ‘weightings’ are largely subjective, and diversity at the scale of biogeographical regions. This multilevel diversity is
consequently diversity indices do not have clear ecological meaning. Such also called inventory diversity.
limitations can be seen mathematically as well. May (1975) pointed out that
diversity indices are simply moments of the full S(N) 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,
distribution, where S is the number of species and N is the total number of 1977) outlined three spatial-levels of differentiation diversity that
individuals. Considering this, he suggested that the variance of the correspond to his inventory diversity: pattern diversity, b diversity, and d diversity.
distribution would be a sensible measure of diversity. He also propounds Of these, b diversity is the most commonly measured ( Mumby, 2001;
that if a diversity index is to be used, then a variance-based index is Vellend, 2001; Crist et al., 2003 ), and it describes change in diversity along
preferable (i.e. one that includes P ð N i= N Þ 2, where N i is the number of a transect or the difference between habitats.
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 3. Biodiversity: what is it and how do we measure it?
attempts to calculate, for a hypothetical individual, the proportion of
encounters that will be with another species ( Hurlbert, 1971 ). 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,
Ni 1998 ), others see it as species diversity ( Bond and Chase, 2002 ), whereas
NK11KX N many propound a much broader definition such as the ‘full variety of life on
Earth’ ( Takacs, 1996 ). NRE (1997) distinguish between native and
where N and N i are as defined above. PIE assumes that each individual introduced species, and others have put extra emphasis on threatened
within a community can encounter or interact with every other individual, an species ( Brockerhoff et al., 2001 ). DeLong (1996) reviewed and assessed a
assumption that would not hold true in reality. Nonetheless, this simple large number of definitions of biodiversity. The International Convention on
index has a clear ecological interpretation. In high PIE communities, there Biological Diversity (2003)
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 uses the following definition:
wind dispersal of pollen ( Hurlbert, 1971 ). It should be noted that Simpson’s
(1949) index is closely related to PIE. “Biological diversity” [biodiversity] means the variability among living
organisms from all sources including,
inter alia, terrestrial, marine and other aquatic ecosystems and the
Describing species diversity as a single value compromises much of the ecological complexes of which they are part; this includes diversity
detailed structure of a community. Theoretical rank-abundance models within species, between species, and of ecosystems.
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 One way to gain an appreciation of the word’s most widely accepted
et al., 1943 ), and MacArthur’s broken stick model ( MacArthur, 1957; Webb, meaning is to consider its evolution. In 1980 the term biological diversity
1974; May, 1975 ). The geometric series and the broken stick describe the was used to describe what was presumably species richness ( Lovejoy
respective opposing extremes where a community is dominated by a few 1980 ). In the same year, Norse and McManus (1980) used this terminology
species and where species are equally abundant. The log series and the to describe a concept that incorporated both ecological diversity and
log normal describe intermediate situations, with the former being closer to genetic diversity. In 1981 the US Strategy Conference on Biological
geometric and latter to the broken stick. A pragmatic limitation of such Diversity was held. In 1985 the
models is that they are not
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 91

( Harper and Hawksworth, 1994 ). Ghilarov (1996) noted that the word impact on the system as a whole ( Walker, 1988 ). There is empirical
biodiversity was originally used in political debate rather than science. But evidence supporting both the diversity-stability ( Tilman and Downing, 1994 )
scientists soon adopted it to secure research funding. In particular, it and species redundancy ( Bellwood et al., 2003 ) hypotheses. Mathematical
seemed to provide a justification for disciplines such as systematics, which reckoning has demonstrated that the stability of any system increases with
traditionally had difficulty convincing funding bodies of the value of their the complexity (i.e. number of components) only until a critical point is
work. This somewhat disingenuous origin in the scientific literature is reached, at which the system becomes unstable ( Gardner and Ashby,
probably responsible for the current ambiguity of the word, especially its 1970; May, 1972 ). Species diversity has often been discussed in relation to
synonymous treatment with ecological diversity, species diversity, and ecosystem functioning. But here we run into a further problem—how do we
species richness. 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
There are many ways of measuring biodiversity. Clearly, the indices contain ( Ghilarov, 2000 ). Furthermore, the relationship between species
and models described earlier are applicable if we consider biodiversity to diversity and ecosystem functioning appears to change over different
be synonymous with species diversity. However, metrics that accommodate spatial scales ( Bond and Chase, 2002; Jonathan, 2002 ).
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 Ghilarov (2000) has argued that more emphasis should be put on the
and proximity to a stand of native vegetation. The Biodiversity Benefits intrinsic worth of biodiversity as a rationale for valuing it. He states that “if
Index (BBI) is an extension/modification of the habitat hectares index ( Oliver biodiversity has intrinsic value it could in principle be ‘useless’ for human
and Parkes, 2003 ). Neither of these metrics have strict ecological meanings needs or for ‘ecosystem functioning’.” This is an important argument, as it
( c.f. Hurlbert’s PIE). The weights assigned to the various components are removes the anthropocentric theme that underlies many justifications for
purely subjective in the sense that they do not represent actual ecological protecting biodiversity. There is a danger that the public or industry will
processes, and the statistical distributions associated with each component have a suspicious opinion of biodiversity if its benefits are mostly expressed
are not considered. Moreover, as noted by Williams (2004), the ecological in terms of reimbursements to humans. Such (perceived) benefits are likely
significance of the mathematical manipulations applied to components, to be indirect or even non-existent, and this could lead to a devaluing or
summation for habitat hectares and multiplication for BBI, used to obtain mistrust of biodiversity. People need to be aware of issues such as the
the metric are not explained or justified. uniqueness of species, the right of species to exist, and the irreversible
nature of extinction.

5. Conclusion
4. Justifications for protecting species diversity or biodiversity
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,
The study of species diversity, or at least species richness, gives on the other hand, is an unclear concept, and its value to the ecologist is
ecologists insights into the stability of communities ( Walker, 1988 ). The questionable. Nonetheless, it is well entrenched in the world of
relationship between species diversity/richness and community stability is environmental management and policy, and it may be that it is a useful tool
quite complex. Stability can be defined as the ability of a system to recover from a sociological and political perspective, even if it does have substantial
to an equilibrium state after disturbance, or simply persistence of the theoretical limitations. Ultimately, if biodiversity plays a key role in
system ( May, 1976 ). The diversity-stability hypothesis asserts that species minimising anthropogenic impacts on nature, then it is indeed valuable as a
vary in their traits, and that in a highly diverse (species rich) system there word and concept.
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 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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