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Research
Projects
My research encompasses a number of fields, but I think the common
theme is an interest in biological endemism. I study species and
groups of species (particularly land snails) that are restricted
to small areas, like individual caves, valleys or hills. This
means some of my studies are taxonomic or biogeographic in focus,
whereas others deal with the ecology and population genetics of
speciation. Besides this there are conservation biological implications
to be studied. Below are brief descripions of the main ongoing
projects. The corresponding research papers can be found in the
list of publications.
The evolution
of short-range endemics on limestone outcrops in Malaysia
Throughout Malaysia, limestone appears at the surface as small,
steep-cliffed hills, on average just half a kilometre in diameter,
and mostly surrounded by alluvial, non-calcareous soils. In total
almost 800 separate hills are known (visit Liz Price's WildAsia
site to learn more on Malaysian karst). Many land snails thrive
on these calcium-rich, alkaline soils. Certain groups, in particular
Diplommatinidae, Streptaxidae, and Vertiginidae, show high degrees
of endemism, with strong differentiation in shell shape from one
hill to the next and many species that are endemic to just a single
hill. Using a combination of population genetics, morphometrics,
and field experimentation, we have been trying to understand the
factors involved in the evolution of these limestone endemics,
in particular within the subgenus Plectostoma of the genus Opisthostoma
in Borneo, but also in the vertiginid Gyliotrachela from West-Malaysia.
The former group is targeted in a system involving some 15 outcrops
along the lower Kinabatangan river valley in eastern Sabah. In
addition to evolutionary studies, we study the value of these
outcrops in land snail conservation. Funding comes from Universiti
Malaysia Sabah, the Treub Foundation, the Clive Marsh Sabah Conservation
Fund, and the Royal Netherlands Embassy in Kuala Lumpur.
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an
isolated limestone outcrop in Sarawak.
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correlation
between genetic and geographic distance among populations
of Gyliotrachela hungerfordiana from West-Malaysian
limestone hills.
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| Gyliotrachela
hungerfordiana |
Opisthostoma
(Plectostoma) mirabile |
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| Two
coiling morphs of A. inversus from West-Malaysia. |
The maintenance
of coil dimorphism in Amphidromus
The tropical tree snails of the genus Amphidromus are well-known
for their colourful shells and their diversity throughout Southern
Asia (see also this
website of Conchologists of America for a nice article on
this snail genus). One of the subgenera, Amphidromus s. str.,
is also famous for being one of the few groups of snails in which
coil dimorphism is common: in most species, both right-handed
and left-handed snails occur together in a population. If, as
is the case in most land snails, mating between individuals of
opposite coil is hampered, there should always be selection against
the rarer morph and the population should quickly evolve towards
coiling monomorphism. However, this does not seem to happen. In
the Bornean species Amphidromus martensi, for example,
all populations show morph frequencies that are close to 50%-50%.
This project aims at understanding what maintains the coil polymorphism,
by mapping spatial distributions of coiling morphs and by observing
copulations in A. martensi and A. inversus. It is
funded by Universiti Malaysia Sabah and the Treub Foundation.
The evolution
of cave snails
Tropical cave snails are good candidates for parapatric speciation
across a steep ecotone. For limestone caves in Borneo, the external
environment is warm, bright, and ecologically very complex, whereas
the cave environment, just a hundred meters from the cave entrance,
is cool, dark, and biologically poor. These different conditions
may force cave snails to adapt in the face of ongoing gene flow.
We study a system of two Georissa snails, a troglobite and its
epigean ancestor, in a limestone cave in the interior of Sabah,
Malaysian Borneo, using morphometrics and population genetics.
The results so far show that the cave snail is indeed descended
from the local population of G. saulau that inhabits the
cliffs of the hill in which the cave is located. This project
is funded by Universiti Malaysia Sabah and the Danish Corporation
for Environmental Development.
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| impressions
of the field site at Gua Sanaron, Sabah |
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| Left:
the epigean Georissa saulae; right: the (undescribed)
descendant cave species; centre: an intermediate form from
the cave entrance. |
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| A Principal
Component Analysis, showing that the epigean G. saulae
(OUT-1 and OUT-2) and the cave species living deep in the
cave (IN-2) have very different shell shapes. The population
that lives in the cave entrance (IN-1) has an intermediate
form. |
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| a phylogeny
based on mtDNA, which shows that the cave species (G. n.
sp.) is closely related to the local population of G. saulae. |
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| daybreak
over Gunung Kinabalu |
The evolution
of high-altitude endemics on Mount Kinabalu
At 4,100 metres high, Mount Kinabalu (click
here for UNEP's data sheet on this World Heritage site) is
one of the few existing, relatively pristine tall tropical mountains,
unique for displaying an uninterrupted series of habitats from
lowland tropical rainforest at the foot to subalpine scrub at
the summit. The numbers of species endemic to the higher elevations
are staggering, especially in taxa like angiosperms, insects and
land snails, but even certain bird and mammal species are restricted
to Mount Kinabalu and its surrounding peaks. In land snails, many
of the high-altitude endemics appear to have existing sister species
at the lower elevations. In some cases, several closely related
species succeed each other along the elevational gradient. Using
detailed mapping of distribution patterns, morphometrics and population
genetics, we aim to understand the evolution of endemics on this
relatively young (1 million years) mountain. Funding is from Universiti
Malaysia Sabah.
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| taking
a break from field work at Panar Laban, at 3,500 m altitude. |
A selfish
symbiont to improve the biocontrol agents of the Cocoa Pod Borer
The Malaysian
Cocoa Board uses several trichogrammatid egg parasitoids (minute
parasitic wasps) to control the Cocoa Pod Borer, a moth that affects
cocoa. The wasps are mass-cultured in their labs in Tawau and
distributed among cocoa farmers for release. We are attempting
to artificially infect these parasitoids with a strain of the
endosymbiotic bacterium Wolbachia to produce an all-female,
parthenogenetic strain that will be more effective at controlling
the pest. This project is funded through the Malaysian government's
Intensifying Research Priority Areas programme.
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