Accelerat ing t he world's research.

Singapore: half full or half empty?

Richard T Corlett

Related papers Download a PDF Pack of t he best relat ed papers 

Dramat ic decline in bat species richness in Singapore, wit h implicat ions for Sout heast Asia
Benjamin P. Y-H. Lee

Cat ast rophic ext inct ions follow deforest at ion in..
Andrea Mart nez

Local geographic range predict s freshwat er fish ext inct ions in Singapore
Heok Hee Ng
CHAPTER 17
Singapore
Half Full or Half Empty?
Richard T. Corlett
Xishuangbanna Tropical Botanical Garden, Chinese Academy of Sciences, Menglun, Yunnan, China
SUMMARY
Singapore has had an inluence on the development of
tropical Asian biology over the past 200 years that is
disproportionate to its size, through the many scientists
who have been based there. Meanwhile, the island
itself was subject to largely uncontrolled deforestation
and exploitation, leaving only small areas of protected
forest and a depleted native biota, while alien species
dominate outside the nature reserves. Yet the biota is
still very rich and new species continue to be found.
Singapore has also proved immensely valuable as a
relatively well-documented case study of extreme
human impacts in the equatorial tropics.
INTRODUCTION
Singapore was once described by the Indonesian Presi-
dent Habibie as a “little red dot” and this epithet has
subsequently been adopted by Singaporeans as a
symbol of the nation’s success, despite limitations of
size (704 km2), population (5.3 million), and natural
resources (virtually none). That size is not everything
is shown by Singapore’s per capita gross domestic
product, which ranks it with the richer countries of the
European Union, and the global top three rankings for
Conservation Biology: Voices from the Tropics, First Edition. Navjot S. Sodhi, Luke Gibson, and Peter H. Raven.
© 2013 John Wiley & Sons, Ltd. Published 2013 by John Wiley & Sons, Ltd.
its port and airport. Less positively, a recent global
review ranked Singapore worst for its environmental
impact in relation to total resource availability (Brad-
shaw, Giam and Sodhi, 2010), although it can be
argued that this is an inevitable consequence of the
late twentieth-century model of development applied
successfully to an island city state.
Biogeographically, in contrast, Singapore is nothing
special, being separated from the southern tip of the
Asian mainland by shallow straits less than a kilometer
wide at the narrowest point. As one would expect, there
are few endemic species and the local diversity in those
groups of organisms for which comparable data are
available is lower than at equivalent sites on the main-
land or on the much larger neighboring islands of
Borneo and Sumatra. Singapore’s disproportionate
inluence on the development of biology in tropical
Asia relects not its own biota, therefore, but the efforts
of the people who have worked there since the modern
settlement was founded by Stamford Rafles in 1819.
NINETEENTH CENTURY: EXPLOITATION
AND DEFORESTATION
Rafles himself was an enthusiastic amateur biologist,
but the best description of Singapore in 1819 comes
 Asia: Singapore 143

from his assistant, the surgeon–botanist William Jack,

in a letter to his family on June 20:

It is impossible to conceive anything more beautiful

than the approach to Singapore, through the archi-
pelago of islands that lie at the extremity of the
Straits of Malacca. Seas of glass wind among innu-
merable islets, clothed in all the luxuriance of tropi-
cal vegetation and basking in the full brilliance of a
tropical sky . . . I have just arrived in time to explore
the woods before they yield to the axe, and have
made many interesting discoveries, particularly
of two new and splendid species of pitcher-plant
[Nepenthes raflesiana and Nepenthes ampullaria], far
surpassing any yet known in Europe.

As Jack foresaw, the primeval forest rapidly did “yield

to the axe,” but nineteenth-century Singapore, with its
busy port, easily accessible habitats, and relatively
comfortable living conditions, continued to attract a
long list of famous – or soon to be famous – biological
visitors and residents (Ng, Corlett and Tan, 2011).
Most famous of all was Alfred Russel Wallace, who
stayed in Singapore several times between 1854 and
1862 during his collecting trips throughout the Malay
Archipelago. His local collections were mostly of beetles
(Figure 17.1):

In about two months, I obtained no less than 700

species of beetles, a large proportion of which were
quite new. . . . Almost all these were collected in one
patch of jungle, not more than a square mile in extent,
and in all my subsequent travels in the East I rarely if
ever met with so productive a spot. (Wallace, 1869)
This “patch of jungle” was Bukit Timah hill, at 164 m
the highest point in Singapore. Wallace attributed the
beetle diversity in a large part “to the labors of the
Chinese wood-cutters” who had “furnished a continual
supply of dry and dead and decaying leaves and bark,
together with abundance of wood and sawdust, for the
nourishment of insects and their larvae.”
Uncontrolled deforestation for cash crops (gambier,
pepper, and others) and exploitation of the remnants
(for timber, rattans, irewood, and game) took its inevi-
table toll. By the time Nathaniel Cantley, Superintend-
ent of the Singapore Botanic Gardens, published his
Report on the Forests of the Straits Settlements, in 1884,
there were serious concerns about the timber supply
(Cantley, 1884). As Cantley reported:
Figure 17.1 Twelve new species of checkered beetles
(Cleridae) collected by Alfred Russel Wallace “within a few
days after his arrival at Singapore” in 1854. Reproduced
from Westwood (1855) Descriptions of some new species of
Cleridae collected at Singapore by Mr. Wallace. Proceedings
of the Zoological Society of London 23, 19–26.
Such Crown forests as remain uncut are widely dis-
tributed in isolated patches over the island. These
forest patches or clumps are of various sizes, from
half an acre or so to about 25 acres [10 ha], and of
no particular shape; their distance from each other
may average a quarter of a mile though often exceed-
ing a mile. The interspace is generally waste grass-
land, which supports, as a rule, only the strongly
growing grass known locally as “lalang” [Imperata
cylindrica].”
144 Conservation Biology
Extinction from isolated forest fragments can be a slow
process and when H. N. Ridley, Cantley’s successor in
charge of the Botanic Gardens, described their fauna
in 1895, they were still “haunted by numerous
mammals” (Ridley, 1895). Ridley describes the scene at
dusk when:
[t]he wild pigs are making onslaughts on the pine-
apple and tapioca ields, the deer come out to crop
the shrubs on the edge of the woods, the tiger is
moving quietly through the bushes in pursuit of the
pigs and deer, . . . the large red lying squirrel is
taking its surprising leaps from one lofty tree to
another, . . . [and] the great fox-bats are lapping
slowly overhead.
All these species were lost from the main island of Sin-
gapore over the coming century (although pigs have
recently reinvaded).
This delay in extinction after the major period of
nineteenth-century forest clearance is important
because, with a few exceptions (such as the botanical
records of Jack and Wallich, and the insects collected
by Wallace), the period from 1870 to 1900, when
>80% of the forest had already gone, marked the start
of serious biological collections and thus the “baseline”
on which later extinction estimates have been based. It
is also important in predicting the ultimate fate of
present-day examples of recently cleared tropical land-
scapes, where the diversity of forest vertebrates can
give a misleading impression of the long-term capacity
of highly fragmented forests to support viable popula-
tions of forest animals.
TWENTIETH CENTURY: EXTINCTIONS,
INVASIONS, AND CONSERVATION
Cantley’s report led to the establishment of a system of
forest reserves in Singapore. These eventually made up
10% of the land area, but they were by no means
entirely forested. In any case, they were not taken seri-
ously enough to apply much of a brake to the contin-
ued erosion of what little forest area remained. The
biodiversity that had attracted so many biologists in the
nineteenth century began to decline. When Ridley pub-
lished his Flora of Singapore in 1900, he commented
that some forest species recorded by Danish botanist
Nathaniel Wallich in 1822 “appear to have quite van-
ished” while many open-country aliens had become
established over the same period (Ridley, 1900). The
extinction of native, mostly forest-dependent, species and
the establishment of alien, open-country species are
recurring themes in natural history publications on Sin-
gapore throughout the twentieth century (Corlett, 1992).
Perhaps ironically, Ridley is most famous as the man
responsible for establishing the rubber industry in
Southeast Asia, and it was largely thanks to him that
the patchwork of habitats described by Cantley in 1884
had been replaced by the 1930s, over 40% of Singa-
pore, by a rubber monoculture. F. N. Chasen, who later
became Director of the Rafles Museum, commented in
the introduction to the irst comprehensive checklist
of the birds of Singapore that “rubber estates are
notoriously unproductive from a naturalist’s point of
view,” and went on to list bird species that had been
lost since the nineteenth century, including all pheas-
ants (although Chasen doubted the validity of the
nineteenth-century records), hornbills, and trogons
(Chasen, 1923). Much of the unprotected forest that
still survived was on swampy ground, but most of this
freshwater swamp forest was cleared in the 1920s and
1930s. We would know nothing of its botanical com-
position were it not for the efforts of E. J. H. Corner,
Assistant Director of the Singapore Botanic Gardens,
who used these fellings as an opportunity to make com-
prehensive collections of the trees, climbers, and epi-
phytes (Corner, 1978).
The fact that any primary forest at all survived to the
present day is largely due to the efforts of Corner’s boss,
the Director of the Gardens, Eric Holttum. In 1936,
when the forest reserves established by Cantley were
abolished, he managed to get Bukit Timah regazetted
(after a brief and damaging period without protection),
“on grounds of amenity . . . and botanical interest”: still
a forest reserve in name, but a nature reserve in prac-
tice. Then, after World War II, Holttum again inter-
vened to stop the increasing threat from encroaching
granite quarries and the resulting enquiry resulted in
the Nature Reserves Act of 1951 and the formal crea-
tion of Bukit Timah Nature Reserve (BTNR). BTNR and
the adjacent Central Catchment Nature Reserve form
the non-coastal component of the present nature
reserve system, protecting all the surviving primary
forest fragments (totaling circa 200 ha) and most of the
native secondary forest (circa 1600 ha). The other
reserves are Sungei Buloh Wetland Reserve (130 ha),
consisting largely of secondary coastal wetlands, and
Labrador Nature Reserve (10 ha), which contains sec-
ondary dry coastal forest and cliff vegetation.

When Navjot Sodhi arrived in Singapore in 1995, he
became interested in the island’s potential as a well-
documented “worst-case scenario” from which general
conservation principles could be derived. He started
with an analysis of bird extinctions (65 more species
had gone since Chasen’s 1923 list) (Castelletta, Sodhi
and Subaraj, 2000). Over the next decade, he and his
students looked at bees (Liow, Sodhi and Elmqvist,
2001), butterlies (Koh, Sodhi and Brook, 2004), dung
beetles (Lee et al., 2009), and angiosperms (Sodhi et al.,
2008). Data on decapods, phasmids, ishes, amphibi-
ans, reptiles, and mammals were added in the classic
paper, “Catastrophic extinctions follow deforestation in
Singapore” (Brook, Sodhi and Ng, 2003), which sum-
marized the lessons learned from Singapore. These
lessons included the high overall extinction rates (28–
73%, depending on assumptions made about extinc-
tions before the irst reliable species records in the
1870s), the great variation between taxonomic groups
(5–43%, for observed extinctions), the greater vulner-
ability of forest than non-forest species (33% versus 7%
for observed extinctions), and the concentration of the
survivors (>50%) in the tiny nature reserves. The paper
then goes on to infer likely extinctions resulting from
deforestation elsewhere in Southeast Asia over the next
century.
TWENTY-FIRST CENTURY SINGAPORE:
HALF FULL OR HALF EMPTY?
The pessimistic (half-empty) case for Singapore biodi-
versity is easy to make. Most of the people mentioned
earlier made their names for work on a larger regional
stage. Singapore was important as a base and, in some
cases, a training ground, but much less so as a study
area. Singapore’s biota was never particularly rich by
regional standards and around half of it has subse-
quently been lost. Moreover, approximately half of the
surviving species in well-studied groups are nationally
endangered (Davison, Ng and Ho, 2008). Deforested
areas were initially occupied by tolerant native plants
and animals, mostly of coastal origin, in the nineteenth
and early twentieth centuries, but these have now been
largely displaced by aliens, which dominate most habi-
tats outside the nature reserves. It is not yet clear if this
alien dominance can ever be reversed, or, conversely,
if it threatens the last areas of native forest (Corlett,
2010). Climate change, in the form of a predicted
3–4°C warming by the end of this century, is an addi-
Asia: Singapore 145
tional threat that we are a long way from being able to
quantify (Corlett, 2011, 2012). The nature reserves
are well protected, but occupy less than 5% of Singa-
pore’s land area and have no marine counterparts, so
the future for Singapore’s native biodiversity does not
look bright.
But there are other ways of looking at Singapore.
First, it is still stunningly rich in native species for a tiny
(704 km2), largely urbanized, island city state. There
are still around 1600 extant species of native vascular
plants, 350 birds, 26 mammals, 117 reptiles, 25
amphibians, 35 freshwater ish, 300 butterlies, 124
dragonlies, and so on (Ng, Corlett and Tan, 2011).
With these numbers, it is not hard to see Singapore as
half full. Moreover, species new to science are discov-
ered almost every month, by both resident and visiting
scientists. More than a hundred such species have been
described in the last few years, including new species
of mosses, fungi, lichens, ishes, nematodes, spiders,
mites, harvestmen, wasps, beetles, bugs, lies, shrimps,
barnacles, and crabs (Ng and Corlett, 2011).
Second, its value as a case study – as recognized by
Navjot Sodhi – has grown with time. Singapore may be
the worst-case scenario for tropical continental biodi-
versity, but the fact that so much has survived – so far
– suggests that extinction is at least slower than we had
feared. Attempts to use Singapore to identify traits that
predict vulnerability to extinction have had varied
degrees of success (Castelletta, Sodhi and Subaraj,
2000; Koh, Sodhi and Brook, 2004; Sodhi et al. 2008),
but this is a useful lesson, suggesting either that we are
investigating the wrong traits, or that extinction can
sometimes be trait-neutral. Moreover, Singapore’s
urbanization has interest beyond its importance as a
driver of extinction. By 2030, the percentage of the
total population that is urban in the Asian, African,
and American tropics is projected to be 52%, 59%, and
84%, respectively (Montgomery, 2008; DeFries et al.,
2010). Indeed, most global population growth is now
in tropical cities and lessons learned in Singapore (e.g.,
on invasive urban birds; Lim et al., 2003) can have wide
application.
Despite these factors that highlight the value of
Singapore’s natural assets, Singapore has been unchar-
acteristically timid when it comes to biodiversity con-
servation, so that many tools in the modern
conservation toolbox remain unused. Aliens have had
a free run outside the nature reserves, ecological resto-
ration has been ad hoc and small scale, and there has
been no systematic attempt to reintroduce species lost
146 Conservation Biology
from Singapore that still survive in the immediately
adjacent areas of Malaysia and Indonesia. A serious
attempt to expand the area of native-dominated forest,
perhaps to 10% of Singapore, to control the most inva-
sive of the aliens, and to reintroduce those species that
succumbed to threats, such as hunting, which have
now been controlled, could transform the situation. We
need to learn how to do these things in the lowland
tropics and Singapore is an excellent place to start.
Climate change may yet overshadow all other threats
to lowland tropical biodiversity, in Singapore and
elsewhere (Corlett, 2011, 2012), but this again is
an opportunity for the world’s only all-equatorial,
all-lowland nation to make a globally signiicant con-
tribution. The close proximity of lowland rainforest
fragments and world-class laboratories makes Singa-
pore a logical center for research into the potential
impacts of climate change. Rigorous, quantitative,
long-term monitoring of tree growth, survival, and
fecundity would be a good start (Clark and Clark,
2011), but the biggest gap in our current knowledge is
information on thermal tolerances and acclimation
capacity needed to predict the future of equatorial bio-
diversity in a warming world (Corlett, 2011).
Finally, the lowland ecosystems of Malaysia and
Indonesia have experienced catastrophic human
impacts over recent decades, with vast areas now
under industrial crop monocultures or expanding
urban areas (Miettinen, Shi and Liew, 2011). It is no
longer safe to assume that a non-endemic species lost
from Singapore still persists elsewhere in the region.
Singapore’s nature reserves are tiny and fragmented,
but they are safe from conversion, logging, and hunting.
If Singapore plays its cards right, it will still be half full
a century from now.
ACKNOWLEDGMENTS
This essay beneited greatly from many years of discus-
sions with Navjot Sodhi, and also with Hugh Tan, Peter
Ng, and many other friends and colleagues in
Singapore.
REFERENCES
Bradshaw, C. J. A., Giam, X. and Sodhi, N. S. (2010) Evaluat-
ing the relative environmental impact of countries. PLoS
One, 5, e10440.
Brook, B. W., Sodhi, N. S. and Ng, P. K. L. (2003) Catastrophic
extinctions follow deforestation in Singapore. Nature, 424,
420–423.
Cantley, N. (1884) Report on the Forests of the Straits Settle-
ment. Singapore Printing Ofice, Singapore.
Castelletta, M., Sodhi, N. S. and Subaraj, R. (2000) Heavy
extinctions of forest-dependent avifauna in Singapore:
lessons for biodiversity conservation in Southeast Asia.
Conservation Biology, 14, 1870–1880.
Chasen, F. N. (1923) An introduction to the birds of Singapore
Island. Singapore Naturalist, 2, 87–112.
Clark, D.A. and Clark, D.B. (2011) Assessing tropical forests’
climatic sensitivities with long-term data. Biotropica, 43,
31–40.
Corlett, R. T. (1992) The ecological transformation of Singa-
pore: 1819–1990. Journal of Biogeography, 19, 411–420.
Corlett, R. T. (2010) Invasive aliens on tropical East Asian
Islands. Biodiversity and Conservation, 19, 411–423.
Corlett, R. T. (2011) Impacts of warming on tropical lowland
rainforests. Trends in Ecology and Evolution, 26, 606–613.
Corlett, R. T. (2012) Climate change in the tropics: the end of
the world as we know it? Biological Conservation, 151,
22–25.
Corner, E. J. H. (1978) The freshwater swamp-forest of South
Johore and Singapore. Gardens’ Bulletin, Singapore, Supple-
ment 1, 1–266.
Davison, G. W. H., Ng, P. K. L. and Ho, H. H. (2008) The Sin-
gapore Red Data Book: Threatened Plants and Animals of Sin-
gapore, 2nd edn. Nature Society, Singapore.
DeFries, R., Rudel, T. K., Uriarte, M. and Hansen, M. (2010)
Deforestation driven by urban population growth and agri-
cultural trade in the twenty-irst century. Nature Geoscience,
3, 178–181.
Koh, L.P., Sodhi, N. S. and Brook, B. W. (2004) Coextinction
of tropical butterlies and their host plants. Biotropica, 36,
272–274.
Lee, J. S. H., Lee, I. Q. W., Lim, S. L.-H., Huijbregts, J. and Sodhi,
N. S. (2009) Changes in dung beetle communities and asso-
ciated dung removal services along a gradient of tropical
forest disturbance in South-East Asia. Journal of Tropical
Ecology, 25, 677–680.
Lim, H. C., Sodhi, N. S., Brook, B. W. and Soh, M. C. K. (2003)
Undesirable aliens: factors determining the distribution of
three invasive bird species in Singapore. Journal of Tropical
Ecology, 19, 685–695.
Liow, L. H., Sodhi, N. S. and Elmqvist, T. (2001) Bee diversity
along a disturbance gradient in tropical lowland forests of
Southeast Asia. Journal of Applied Ecology, 38, 180–192.
Miettinen, J., Shi, C. and Liew, S. C. (2011) Deforestation rates
in insular Southeast Asia between 2000 and 2010. Global
Change Biology, 17, 2261–2270.
Montgomery, M. R. (2008) The urban transformation of the
developing world. Science, 19, 761–764.
Ng, P. K. L. and Corlett, R. T. (2011) Biodiversity in Singapore:
an overview, in Singapore Biodiversity: An Encyclopedia of the

Natural Environment and Sustainable Development (eds P. K. L.
Ng, R. T. Corlett and H. T. W. Tan), Editions Didier Millet,
Singapore, pp. 18–36.
Ng, P. K. L., Corlett, R. T. and Tan, H. T. W. (eds) (2011) Singa-
pore Biodiversity: An Encyclopedia of the Natural Environment
and Sustainable Development. Editions Didier Millet, Singapore.
Ridley, H. N. (1895) The mammals of the Malay Peninsula.
Natural Science, 6, 23–29, 89–96, 161–166.
Ridley, H. N. (1900) Flora of Singapore. Journal of the Straits
Branch of the Royal Asiatic Society, 33, 27–196.
Asia: Singapore 147
Sodhi, N. S., Koh, L. P., Peh, K. S.-H., Tan, H. T. W., Chazdon,
R. L., Corlett, R. T., Lee, T. M., Colwell, R. K., Brook, B. W.,
Sekercioglu, C. H. and Bradshaw, C. J. A. (2008) Correlates
of extinction proneness in tropical angiosperms. Diversity
and Distributions, 14, 1–10.
Wallace, A. R. (1869) The Malay Archipelago. Macmillan,
London, pp. 37–38.
Westwood, J. O. (1855) Descriptions of some new species of
Cleridae collected at Singapore by Mr. Wallace. Proceedings
of the Zoological Society of London, 23, 19–26.