Content uploaded by Nimal Gunatilleke
Author content
All content in this area was uploaded by Nimal Gunatilleke on Oct 17, 2017
Content may be subject to copyright.
alia, terrestrial, marine, and other aquatic ecosystems,
and the ecological complexes of which they are a part;
this includes diversity within species, between species
and of ecosystems’. These three levels of biodiversity,
proceeding from the most obvious (ecosystem and
species diversity) to the relatively less obvious genetic
or within-species diversity may be defined as follows:
Ecosystem Diversity: The sum total of all interrelated
and interacting organisms and all the components of the
physical environment (climate, soil, water, fire, rocks,
wind, landscapes etc.) that function together as one unit.
Depending on the organisms and environment that make
up an ecosystem, different types of ecosystems may be
identified. It is this variability among ecosystems that is
referred to as ecosystem diversity.
Species Diversity: The variation among species is
recognized as species diversity.
Genetic Diversity: The genetic variation among
individuals of a population and that among all the
different populations of the same species comprises the
total genetic variation of that species.
This paper is presented in three parts. Part one,
which includes a substantial component of the paper,
identifies Sri Lanka’s biodiversity at the ecosystem,
species and genetic levels. At the ecosystem level, each
major ecosystem is introduced by giving its distribution,
vegetation stature where relevant, dominant plant taxa,
endemic component, some information on the fauna and
finally, a few references to sites studied for that ecosystem.
At the species level, the number of taxa in each group,
the proportions of endemics and threatened species are
highlighted. At the genetic level, genetic variation within
and between populations of some indigenous species for
which data are available are given along with a brief
overview of genetic diversity in cultivated species.
Abstract : This paper reviews the information available on
the biodiversity of Sri Lanka, the causes of its degradation
and the policies, legal instruments and strategies developed
both nationally and internationally for its conservation and
sustainable use.
In the first part of this paper, the biodiversity of Sri Lanka
at the ecosystem, species and genetic levels is reviewed. At
ecosystem level, each major ecosystem is introduced by giving
its distribution, vegetation stature where relevant, dominant
plant taxa, endemic component, some information on the
fauna and finally, a few references to sites studied for that
ecosystem. At the species level, the number of taxa in each
group, the proportions of endemics and threatened species are
highlighted. At the genetic level, genetic variation within and
between populations of some indigenous species for which
data are available are given, along with a brief overview of
genetic diversity in cultivated species.
The second part examines causes of loss of biodiversity at the
ecosystem level; the third presents information on the policies,
legislation and national strategies related to biodiversity
conservation in the country and in meeting the international
obligations for conservation, sustainable development and
sharing of benefits arising from conservation. A strong plea is
made on both formal and non-formal education at all levels for
using the protected area network as learning laboratories for
testing and demonstrating innovative approaches to reconcile
the conservation of biodiversity with social and economic
development in line with the Man and the Biosphere concept.
Keywords: Conservation, ecosystems, endemics, endemism,
species.
INTRODUCTION
Biodiversity in simple terms is the variety of life on
earth at all levels of biological organization. The legally
accepted definition adopted by the United Nations
Convention on Biological Diversity is ‘the variability
among living organisms from all sources, including, inter
Biodiversity of Sri Lanka
Nimal Gunatilleke1*, Rohan Pethiyagoda2 and Savitri Gunatilleke1
1Department of Botany University of Peradeniya, Peradeniya.
2Wildlife Heritage Trust of Sri Lanka, P.O. Box 66, Mt. Lavinia.
J.Natn.Sci.Foundation Sri Lanka 2008 36 Special Issue 25-62
*Corresponding author
Nimal Gunatilleke et. al
26
Journal of the National Science Foundation of Sri Lanka 36 Special Issue
November 2008
The second part examines causes of loss of biodiversity
at the ecosystem level; the third presents information
on the policies and legislation related to biodiversity
conservation in the country and international obligations
for conservation, sustainable development and sharing
of benefits arising from conservation.
Ecosystem diversity of Sri Lanka
Despite its small size, Sri Lanka has a rich
ecosystem diversity because of its topographic and
climatic heterogeneity as well as its coastal influence.
These natural/semi-natural ecosystems are broadly i) the
marine and maritime or coastal ecosystems, ii) natural
forest ecosystems, iii) natural and semi-natural grassland
ecosystems, and iv) the inland wetland ecosystems.
Table 1 lists the diversity of the different ecosytems
and their extents, where this information is available.
In addition, there are other man-made ecosystems such
as agro-ecosystems (including home gardens) that
structurally resemble natural forests, and cash-crop fields
and plantations.
The following ecosystems in Sri Lanka have
been internationally recognized for their outstanding
biological wealth and their current conservation status.
i. Sinharaja lowland and lower montane rain forest is a
Natural World Heritage Site and also an International
Man and the Biosphere Reserve.
ii. Hurulu dry mixed evergreen forest and the Kanneliya
-Dediyagala - Nakiyadeniya lowland rainforest
complex are International Man and the Biosphere
Reserves.
iii. Western Ghats and Sri Lanka together form one of
the 34 Biodiversity Hotspots1.
iv. Bundala, Anavillundawa and Maduganga estuary
and mangrove ecosystem are Ramsar Wetland Sites.
v. 41 of the 83 wetland sites in Sri Lanka are included
in the Asian Directory of Wetland Sites2.
The marine ecosystems are represented by coral
reefs and sea grass beds and the ocean that surrounds
Sri Lanka (Figure 1).
Coral reefs, both near-shore reefs (mostly fringing,
sandstone, rock and boulder) and well-developed
off-shore reefs are found along 2-3 % of the island’s
coastline. They have a rich diversity of corals with
about 183 species from 68 genera, principally Acropora,
Montipora, Porites, Favia, Favites, Pocillopora,
Goniastrea, Platygyra and Leptoria. Green, red, and
brown algae, mollusks, sea cucumbers, sea anemones,
lobsters and reef fish abound in these reefs. It is estimated
that about 1000 species of fish inhabit Sri Lanka’s coral
reefs3.
Sea-grass beds are found in calm shallow seas,
estuaries and lagoons around the island where 12 species
Coastal Ecosystems
1. Coral reefs not available
2. Sea grass beds 23,819
3. Salt marshes 33,573
4. Mangroves 12,189
5. Sea shores/beeches not available
6. Mud flats not available
7. Lagoons and estuaries 158,017
8. Sand dunes 7,606
Inland Aquatic Systems
1. Fresh water marshes 10,000**
2. Rivers and streams
Riverine Forests 22,435
3. Reservoirs* 170,000
Natural forest ecosystems
1. Tropical lowland wet evergreen
forests or lowland rain forests 141,506
2. Tropical submontane forests 68,616
3. Tropical montane forests 243,886
4. Tropical moist evergreen forests 1,090,981
5. Tropical dry mixed evergreen forests 464,076
6. Thorn scrub forests not available
Natural grassland ecosystems
1. Wet patanas not available
2. Dry Patanas* 65,000
3. Savannas* not available
4. Thalawas* not available
5. Damanas* 10,000
6. Villu
Present
extent (ha)
Aquatic ecosystem
diversity
Present
extent (ha)
Terrestrial ecosystem
diversity
* Man-influenced ecosystems. ** Also includes the villus.
Table 1: Ecosystem diversity and their extents in Sri Lanka
Biodiversity of Sri Lanka 27
Journal of the National Science Foundation of Sri Lanka 36 Special Issue November 2008
Figure 1 : Hikkaduwa marine sanctuary with near-shore coral reef communities. These together with sandy beaches and
the fishing harbour have increased the anthropogenic pressure on this fragile maritime natural resource.
Figure 2 : A mangrove formation dominated by Rhizophora spp. fringing the Negombo Lagoon. Inset: a flower and a fruit of
Sonneratia caseolaris.
(a) (b)
Nimal Gunatilleke et. al
28
Journal of the National Science Foundation of Sri Lanka 36 Special Issue
November 2008
Figure 4 : (a) Canopy of the lowland rain forest of Sinharaja World Heritage Site dominated by Doona (Shorea) and Mesua species exhibiting
habitat specialization leading to greater species richness. (b) A vertical profile as seen in a forest fragment near Kukule Ganga reservoir.
(a) (b)
Figure 3 : Sea shore vegetation dominated by a) Spinifex littoreus which is being invaded by the prickly pear b) Opuntia sp., an exotic xerophyte.
In the background are the woody elements of the sea-shore vegetation dominated by Pandanus sp. which along with other shrubby
vegetation acts as a barrier to coastal erosion and even tsunami.
(a) (b)
Biodiversity of Sri Lanka 29
Journal of the National Science Foundation of Sri Lanka 36 Special Issue November 2008
of angiospermous halophytes have been recorded so far.
Amongst them, Enhalus acoroides, Thalassia hemprichii,
Halophylla and Najas are typical of this ecosystem.
These species have rhizomatous stems and thin •oating
leaves ranging widely in size and shape among the
taxa4,5. They support a diversity of organisms, including
!sh that directly feed on sea grass, while others such as
peneid shrimp larvae feed on grass detritus. Sea-grass
beds also provide habitats for sea turtles and dugong3.
The near-shore and off-shore waters of the Indian Ocean
surrounding Sri Lanka are inhabited by predatory !sh
such as reef sharks, and a range of food and shell-!sh,
lobsters, upon which the marine !shery is dependent. In
addition, there are about 38 species of marine mammals
comprising one locally depleted species of dugong and
37 cetaceans including the sperm whale, the blue whale
and over 20 species of dolphins3.
a) The maritime ecosystems:
These are in•uenced by their proximity to the sea and
include the salt marshes, mangroves and seashore or
strand vegetation.
The salt marshes are con!ned to the inter-tidal •ats
frequently inundated with salt water in the northwestern
and southeastern arid zones of Sri Lanka, covering an
estimated area of 34,000 ha3, 4. They are dominated by
the halophytic or salt-loving plants Halosarcia indicum,
Salicornia brachiata, Sueda maritima, and S. monoica,
which are con!ned to this ecosystem. In addition, there
are about 17 species of salt-marsh associates, which may
be found elsewhere as well5, 6. Marshes also provide
habitats for waterfowl and milk!sh. On inundated higher
ground the salt marshes give way to a distinctive, sandy
seashore vegetation (see below).
The mangroves, in•uenced by tidal amplitude, scattered,
and restricted to coastal lagoons and estuaries; they are
best seen near Jaffna, Batticaloa, Kalpitiya, Rekawa
and Trincomalee (Figure 2). Mangroves are dominated
by the plant families Rhizophoraceae, Acanthaceae
and Avicenniaceae and the tree species Rhizophora
apiculata, Rhizophora mucronata, Ceriops tagel,
Bruguiera gymnorhiza, Acanthus illicifolius, Lumnitzera
racemosa, Avicennia officinalis and Avicennia marina.
Altogether, 18 true mangrove species have been
described from Sri Lanka7. All these species are
adapted to grow under saline conditions and a •uctuating
water table. Nypa fruticans, Sonneratia caseolaris,
R. apiculata and Bruguiera sexangula have been reported
from riverine mangals, while R. mucronata, C. tagal,
S. alba and A. marina dominate fringing mangals in the
dry and intermediate zones8,9. Other studies have been
done on mangroves in the west coast9-13, the northern part
of the country14 and in relation to their conservation10-15.
Mangroves, while harbouring a rich migrant (both
from inland and sea) and resident faunal component 9,15-20
including !n!sh, shell!sh, clams, crabs, oysters and
shrimp, also provide nursery grounds for numerous
marine organisms and roosting sites for birds 16-20.
The sea-shore vegetation is seen above the high-water
mark on sandy, gently sloping beaches. The vegetation
grades from a very short-statured creeping form near
the seafront, through herbaceous erect shrubs, to a
10–15 m tall littoral woodland (Figure 3). The dominant
vegetation is Ipomea pescaprae in the wet zone of the
country and the grass Spinifex littoreus in the drier parts.
The common herbaceous species here include those of
Vernonia, Hedyotis, Phyllanthus and Crotalaria; the
shrubs include those of Scaevola, Clerodendrum, Morinda
and Calotropis; and the tree component comprises
Pandanus spp., Barringtonia asiatica, Calophyllum
inophyllum and Thespesia populnea. These species have
special adaptations to root in unconsolidated sandy soil,
and to withstand strong winds laden with salt spray
and saline soil-moisture conditions. The seeds of most
species are also adapted to wind or water dispersal.
Sri Lanka has a coastline of 1,585 km, and many
of the beaches fringing the island are wide and sandy,
providing ideal habitats for nesting turtles. Five species of
turtles Caretta caretta, (Loggerhead turtle), Lepidochelys
olivacea (Olive Ridley turtle), Eretmochelys imbricata
(Hawksbill turtle), Chelonia mydas (Green turtle) and
Dermochelys coriacea (Leatherback turtle) use this
habitat for nesting21.
b) The inland forest ecosystems:
They range from wet (aseasonal) zone rainforests to
thorn scrub in the arid zone. Between these extremes
are forests in the seasonally dry and intermediate zones.
Based on the patterns of distribution of the angiosperm
•ora and their endemicity, 15 different •oristic regions
have been recognized in the island22. The diversity of
forest ecosystems and their extents are given in Table 1,
and salient features of each are described below.
Tropical lowland wet evergreen forests or lowland rain
forests, restricted to the southwest of the island, up to
a mean elevation of 900 m, and are at present highly
fragmented23. Sinharaja, the largest block of relatively
undisturbed forest and the selectively logged Kanneliya-
Dediyagala-Nakiadeniya group of forests, best exemplify
Nimal Gunatilleke et. al
30
Journal of the National Science Foundation of Sri Lanka 36 Special Issue
November 2008
them (Figure 4). Reaching 30-45 m in height, they are
dominated by the families Dipterocarpaceae, Clusiaceae,
Sapotaceae, Bombacaceae and Myrtaceae. At the lower
elevations (<100 m), Dipterocarpus zeylanicus and
D. hispidus, and at higher elevations (>100 m) Mesua
ferrea and Shorea trapezifolia, dominate the canopy of
these forests. In the sub-canopy, the common species
are Cullenia rosayroana, C. zeylanica and Myristica
dactyloides, and in the understorey tree layer Xylopia
championii and Garcinia hermonii. Large leaves, drip
tips on leaves, buttresses, cauli•ory, drooping young
twigs, colourful young leaves and fewer lianas and
epiphytes characterize its vegetation.
In these forests as much as 60 – 75 % of the tree
species are endemic to Sri Lanka. Most of their tree
species are soft and medium hardwoods and were logged
primarily for plywood manufacture until a moratorium
on logging was imposed by the government in 1989.
Two of the most valuable hardwood species in these
lowland rainforests are Calamander (variegated ebony,
Diospyros quaesita) and in wet zone •ood-plain forests,
Nedun (Pericopsis mooniana).
In this ecosystem, various aspects of the vegetation,
mostly trees and selected faunal groups have been
studied. These forests are the most diverse and the
richest in endemic biota among terrestrial ecosystems24,
25. Some among the vegetation studied include those at
Sinharaja26, Kanneliya25,27,28, Kottawa25,27, Gilimale25,27,
Delwala29 and almost all the forests >200 ha30.
Spacio-temporal dynamics of the Sinharaja forest is
being studied using a 25 ha Forest Dynamics Study Plot
in Sinharaja31. These studies have revealed that most
tree species in them are specialists and exhibit habitat
associations in relation to the topographic variation of
the land form.
The vertebrate fauna in lowland rainforests have
been studied better than most invertebrate groups
and noteworthy among them are studies on the small
mammals32, avifauna33--35, reptiles36, !shes37-39 and
amphibians40-42.
The bird •ocks recorded in Sinharaja are among the
largest reported in terms of numbers per •ock containing
many endemic and threatened species34-35.
Although some information of the invertebrate
fauna in lowland rain forests has been reported, there
is a great dearth of information or none at all with
respect to other taxonomic groups. Studies among the
invertebrate groups include those on butter•ies43,44,44a,
spiders45, ants46,47, fresh water crabs48 and land snails49,50.
The monotypic endemic relict ant Aneuretus simoni is
also found in this ecosystem.
The lower montane forests are con!ned to middle
elevations (900–1500 m) of the Peak Wilderness,
Knuckles (Dumbara hills), Namunukula and the
Rakwana-Deniyaya ranges (Figure 5). Their canopies are
about 20–25 m high, dominated by the families
Dipterocarpaceae, Clusiaceae and Myrtaceae, with
the following soft and medium hardwood species
being dominant: Shorea gardneri, Calophyllum spp.
Cryptocarya wightiana, Myristica dactyloides and
Syzygium spp. The proportion of endemic tree species here
is about 50%51-55. The herbaceous, shrub and epiphytic
•ora have not been studied in depth: preliminary surveys
however, indicate that they may represent a higher
proportion of endemics in this ecosystem. Most species
of the endemic genus Stemonoporus show a localized
distribution in these montane forests22,56,57.
The Knuckles (Dumbara Hills) range shows much
heterogeneity in its vegetation distribution in relation to
the monsoonal climatic regimes to which it is exposed.
The NE •anks are seasonally somewhat drier during the
SW monsoonal period as compared with the SW-facing
slopes which are more moist throughout the year.
These lower montane forests have not been
investigated extensively, even though some of the richest
habitats are represented in this ecosystem; among them,
Floristic Region 9 the foothills of Peak Wilderness,
spanning from Maskeliya and the Kelani valley in the
northwest, through Ambagamuwa around Kitulgala
in the south and eastwards to Maratenna above
Balangoda- encompasses a centre of exceptionally high
endemism22.
Tree vegetation of the lower montane forests studied
include parts of the Knuckles, Eastern Sinharaja26,55 and
different locations in the Peak Wilderness51-54.
Studies on the fauna in these ecosystems are
discussed, along with those in the montane forests, in the
next section.
The montane forests, restricted to the uppermost
elevations of the country (> 1500 m), are dominated by
the Clusiaceae, Myrtaceae, Lauraceae, Symplocaceae
and Rubiaceae. The common tree species in them
are Calophyllum walkeri, C. trapezifolium, Syzygium
revolutum, S. rotundifolium, S. umbrosum, Symplocos
cochinsinensis, Neolitsea fuscata, Cinnamomum
Biodiversity of Sri Lanka 31
Journal of the National Science Foundation of Sri Lanka 36 Special Issue November 2008
ovalifolium, Litsea ovalifolia and Actinodaphne speciosa.
In most areas, the understorey of the forest is dense and
dominated by species of Strobilanthes, bamboo and
Coleus. About 10 m tall, short-statured trees with gnarled
twisted branches, an abundance of epiphytes dominated
by mosses, leafy liverworts, ferns and orchids, colourful
young foliage, relatively small, thick, leathery leaves
and absence of drip tips are distinguishing features of
its vegetation (Figure 6). With increase in elevation
and windy conditions the canopy species become quite
stunted giving way to pygmy forests.
The bryophyte •ora of island is the richest in the
lower montane and montane ecosystems, but relatively
under-explored59,60.
In this ecosystem studies on phytosociology have
been carried out in the Horton Plains61, Hakgala Strict
Nature Reserve62, and the Knuckles63 forest die back in
Horton Plains64-66, Pidurutalagala67 and Kobonilgala in
Knuckles68.
These rainforest ecosystems too, harbour a
substantial proportion of endemic species of
amphibians41,42, freshwater crabs48 and land snails49,50.
The faunal diversity in the lower montane and
montane forests is quite outstanding. Some taxa are
con!ned or restricted to one or few locations either in
the Rakwana-Deniyaya hills, Central Highlands or the
Knuckles. A few examples of such localized distributions
are exempli!ed by Ceratophora spp.69 (C. erdeleni and
C. karu in Eastern Sinharaja; C. tennenti in Knuckles),
the direct-developing shrub frogs, Philautus spp.70,71
(P. schmarda and P. variabilis Central Highlands only;
P. macropus and P steineri from Knuckles only; P.
lunatus and P. poppiae from Handapan Ella Plains
only), the freshwater shrimp Lancaris spp.72 (L.
singhalensis from only Horton Plains and Nuwara
Eliya; and L. kumariae near Rozella), and the fresh
water crabs73 (Ceylonthelphusa diva from Knuckles; C.
savitriae and Perbrinckia rosae from Morningside; and
P. gabadagei from Peak Wilderness). Other studies
carried out at Knuckles on the herpetofauna, agamid
lizards in relation to disturbance, amphibians and bird
•ocks have also been published74-76.
Tropical moist evergreen forests, about 25-30 m tall,
represent the transition or ecotone between forests in the
aseasonal and seasonal climates. They bear similarities
in species composition to both the tropical lowland wet
evergreen forests and the tropical dry mixed evergreen
forests and some species of their own. Fragments of this
forest type are seen at Barigoda near Kurunegala and
Daragoda near Moneragala, and more extensive areas in
Randenigala and Samanalawewa. The dominant families
here are Anacardiaceae, Sapindaceae, Euphorbiaceae
and Moraceae. The dominant species are Mangifera
zeylanica, Canarium zeylanicum, Filicium decipiens,
Dimorcarpus longan, Nothopegia beddomei and
Gironniera parvifolia. Lianas are abundant in this forest
type. Only about 17 % of the tree species in them are
endemic to Sri Lanka. One of these endemics is Hopea
brevipetiolaris77 on the crest of Doluwakanda, which is its
only habitat. A few of the characteristic elements of this
ecosystem are Tetrameles nudiflora, Antiaris toxicaria,
Artocarpus gomezianus and Pterygota thwaitesii25.
In tropical moist evergreen forest ecosystems,
the vegetation has been studied extensively in the
Victoria-Randenigala-Rantambe (VRR) Sanctuary78,
Samanalawewa79, Barigoda forest25,27 near Kuliyapitiya,
Daragoda forest25,27 near Moneragala, Wasgomuwa
National Park80-82, Kahalla83 and Maduru Oya Forest
Reserve84.
Tropical dry mixed evergreen forests represent
54 % of the island’s natural forest cover and 16 % of
its total land area. Typical examples of them are seen
in the North, Eastern, North Central, North-Western and
Southern Provinces. In the best stands they are about
25 m tall, but shorter in those towards the arid zone
(Figure 7). The families Euphorbiaceae, Sapindaceae,
Ebenaceae, Sapotaceae, and Rutaceae dominate them.
The dominant canopy species are Manilkara hexandra,
Chloroxylon swietenia, Schleichera oleosa, and
Pleurostylia opposita, while those in the understorey tree
layer are Pterospermum suberifolium, Drypetes sepiaria
and Dimorphocalyx glabellus. They comprise a mixture
of evergreen and deciduous canopy species. In general,
the latter are found in the canopy, shedding their leaves
during the late dry period, allowing much light to !lter
to the forest •oor. This enables grass and herbaceous
species to •ourish at the onset of the rainy season
before new foliage on denuded canopy trees emerges.
The appearance of these forests differs seasonally and
depicts an annual cyclical variation in •oristic diversity
of the ground vegetation. Leaves of plant species here
are small, without drip tips; many have compound
leaves. The tree trunks branch lower down and show no
buttresses, compared with those in rainforests. Epiphytes
and lianas are relatively sparse in this forest type.
A signi!cant proportion of this •ora is similar to
that of India, and only about 13 % of the tree species are
endemic to Sri Lanka. These forests are rich in quality
Nimal Gunatilleke et. al
32
Journal of the National Science Foundation of Sri Lanka 36 Special Issue
November 2008
Figure 5 : Lower montane rain forests a) in the Knuckles (Dumbara) conservation forest. These windswept hence low-statured forests are
dominated by Syzigium species (in pink •ush). b) Lower montane forests in eastern Sinharaja dominated by Doona (Shorea) gardneri.
Figure 6 : a and b) Wind-swept and short-starured montane cloud forests interspersed with wet patana grasslands in a mosaic. c) Montane forest
trees with characteristic gnarled tree crowns
(a)
(a)
(b)
(b)
(c)
Biodiversity of Sri Lanka 33
Journal of the National Science Foundation of Sri Lanka 36 Special Issue November 2008
hardwood species suitable for sawn timber and for this
reason and for agricultural expansion, they have been
clear-felled in the past and are now mostly restricted to
protected areas.
In tropical dry mixed evergreen forest
ecosystems, vegetation surveys have been done at
Kahalla83, Ritigala25,27,85,86, Maduru Oya Reserve84,
Polonnaruwa 89,90, Wasgamuwa80-82, Irrigation area of the
Lower Walawe Basin91, and Wilpattu92.
Thorn scrub forests found in the arid zone are typical
of the vegetation in Yala in the southeast and Mannar
region in the Northern Province. As the name implies,
thorny species are abundant here and reach to about 5 m
in height showing no species stratification. Its dominant
families are Salvadoraceae, Mimosaceae, Euphorbiaceae
and Rhamnaceae. The dominant species are Salvadora
persica, Acacia planifrons, Dichrostachys cinerea,
Bauhinia racemosa, Eugenia bracteata, Phyllanthus
polyphyllus, and Zizyphus oenoplia. Endemic plant
species are almost absent in these forests.
In Yala93,94 and Bundala95, this ecosystem together
with its neighbouring grasslands and water bodies,
harbour large mammals (leopard, elephants, spotted
deer, sambhur and wild pig, primates etc.), numerous
water-and inland bird species (both migrant and resident)
reptiles, amphibians and a diversity of invertebrate
species.
c) Natural / Semi Natural Grasslands :
Very broadly, based on elevation, climate, plant
composition and degree of disturbance, six types of
grassland ecosystems are recognized in Sri Lanka96,97
(Table 1). They are the following:
The wet patana grasslands, found around and above
1800 m altitude, are best seen in Horton Plains and
around Nuwara Eliya (Figure 8). They are dominated
by the tussock grass species Chrysopogon nodulibarbis
and Arundinella villosa on higher ground and Garnotia
exaristata and the dwarf bamboo, Sinarundinaria
densifolia, close to waterways. A diversity of delicate,
small herbaceous species with colourful flowers grow
among the grasses. They include Pedicularis zeylanica,
Satyrium nepalense, Exacum walkeri, Osbeckia spp.,
Ranunculus spp. (some very seasonal) and the fern
Pteridium revolutum. Rhododendron arboreum ssp.
zeylanica about 2 m tall, often displaying spectacular
brilliant red flowers, the shorter Gaultheria leschenaultii
whose leaves smell of wintergreen and the invasive spiny
Ulex europaeus, are the widely scattered woody species
in these grasslands (Figure 9).
During less wet periods of the year, February to
April, these grasslands are subject to accidental fires.
These fires, ground frost in January/February, a high water
table and herbivory all contribute to the maintenance of
these grasslands.
In the early 1960s about 120 ha of wet patana
grassland were used for cultivation of seed potato by
the Department of Agriculture and later abandoned,
following which the area was gradually converted to a
carpet-like grassland dominated by the exotic Penesetum
sp. This grassland, with its vegetation growing close to
the ground, does not provide environmental conditions
and microhabitats for the herbaceous flora to thrive,
unlike among the tussock grasses of the natural wet
patana. Therefore, its floristic richness is much less, and
consequently its faunal diversity as well.
Sambhur, wild boar and black-naped hare feed in
these grasslands at night but rest in the adjacent forests
during the day. The grassland/forest complex provides
them an ideal system for survival. Among the other
animals recorded98 are three endemic freshwater crabs
(Ceylonthelphusa scansor, Perbrinckia punctata, P.
glabra), the endemic shrimp (Lancaris sinhalensis)
and among the amphibians Philautus microtympanum,
Fejervaryia greenii, Microhyla zeylanica and
Polypedates eques inhabit the streams and/or their
immediate environs.
Among the wet patana grasslands, the flora and
fauna has been best studied in Horton Plains98-100.
The dry patana grasslands, located between 500 –
1000 m elevation, are relatively widespread, occurring
in the Uva basin, and around Gampola, Nawalapitiya,
Hantana and Rakwana. These grasslands have resulted
from the removal of tree vegetation for agriculture and
subsequent abandonment (Figure 10). The dominant
grass species in them are Cymbopogon nardus and
Themeda tremula, both forming tussocks. Other species
seen among the grasses include a large number of tall
composite herbs (Blumea spp., Vernonia spp. etc.) and
semi-woody to woody shrubs (Cassia spp., Crotalaria
spp., Lantana camara, Osbeckia octandra, Psidium
guajava, Wikstroemia indica).
During the dry months these dry patanas are
purposely burnt to obtain fresh grass for fodder or to hunt
animals. Their very existence is a result of fire. Much of
these grasslands are now converted to agricultural land
and Pinus and Eucalyptus plantations. They are also
being taken over by the aggressively competitive exotic
fodder grass Panicum maximum and invasive species
Nimal Gunatilleke et. al
34
Journal of the National Science Foundation of Sri Lanka 36 Special Issue
November 2008
Figure 7 : Dry mixed evergreen forest along a small hill in Giritale during a) the wet and b) dry season.
Figure 8 : Montane wet patana grasslands cleared and terraced for seed potato farming in the 1970s and later abandoned. The
carpet-like grasses introduced along with seed –potatoes are frequently grazed by sambar. Native tussocky grasses
(a) (b)
Biodiversity of Sri Lanka 35
Journal of the National Science Foundation of Sri Lanka 36 Special Issue November 2008
Figure 10 : The dry patana grasslands in the North-eastern Knuckles range has resulted from clearing the forest vegetation for
planting tea (Camellia sinensis) and later abandonment. These grasslands in the Central and Uva province have been
subsequently planted with exotic tree crops such as Pinus and Eucalyptus spp.
Figure 9 : Rhododendron arboreum var. zeylanicum with showy
•owers is a characteristic tree more abundant in the grassland
than in the neighbouring forest. The dwarf bamboo,
Sinarundinaria densifolia and the fern, Pteridium aquilinium
are also abundant in the grasslands.
Nimal Gunatilleke et. al
36
Journal of the National Science Foundation of Sri Lanka 36 Special Issue
November 2008
like Clusia rosea. Ecology of the vegetation in the dry
patana grasslands has been studied at Hantana101-103.
The savannas occur in the intermediate and dry zones,
between 300–1000 m elevation. Those on the eastern
slopes of the central massif and around the Uva basin
are called upland savannas while those in Moneragala
and Bibile and in the Gal Oya basin are known as
lowland savannas. They resemble a parkland, with
scattered, !re-tolerant trees dominated by Careya
arborea, Phyllanthus emblica, Terminalia chebula,
T. bellirica, all with medicinal properties. The ground
layer is dominated by the grass species Cymbopogon
polyneuros in association with Themeda triandra at
higher elevations and Aristida setacea and Panicum
sp. at the lower elevations. Savannas are maintained by
anthropogenic !res, mostly to facilitate the collection of
fallen fruits of the medicinal species.
These savannas are used as foraging habitats by
elephants and other herbivores, especially when there is
new growth of the grass species after they have been set
on !re. The damanas given below are foraged by these
herbivores.
Savanna ecosystems and neighboring habitats have
been studied in the VRR sanctuary77, 104-107 and in the Gal
Oya catchment108.
Damanas, also known as dry low country parklands or
grasslands, occur in the Ampara and Batticoloa districts
of the eastern dry zone109. They are dominated by the
tussock grass Cymbopogon nardus. Other less abundant
grass species include Aristida setacea, Imperata
cylindrica, Themeda spp., Brachiaria spp., Cyanodon
dactylon, Echinocloa colona, Eragrostis sp. and
Sporobolus diander. Scattered in the grass are herbaceous
(Alisicarpus vaginalis, Desmodium heterophyllum, D.
triflorum), woody shrubs (Calotropis gigantea, Cassia
auriculata, Lantana camara) and trees (Manilkara
hexandra, Pterocarpus marsupium, Acacia leucophloea,
Butea monosperma).
Damanas provide fodder to wild elephants and
buffaloes. Indeed, these animals are largely responsible
for the maintenance of this vegetation, which except
for their impact, is likely to contain a higher density of
woody species. Leaves of Diospyros melanoxylon, locally
known as Kadumberiya, growing in this ecosystem are
widely used as wrappers for local cigarettes (beedi).
Talawa grasslands occur in the Haldumulla area, where
the topsoil is eroded and truncated to some degree97,109.
While the tall (1.5 m) tussock grass Cymbopogon nardus
dominates them, other less abundant grasses seen are
Andropogon lividus, Arundinella villosa, Chrysopogon
aciculatus and Themeda tremula. The herbs and shrubs
scattered among the grass include Desmodium triflorum,
Elephantopus scaber, Evolvulus alsinoides, Lantana
camara and Psidium guava.
Villus are wetland ecosystems also known as freshwater
riverine marshes found in the "oodplains of the
Mahaweli River in the eastern dry zone and in moist
depressions with a perennially high "uctuating water
table (Figure 11). Flooding and rain water accumulation
suppress any tree growth within the villus but encourage
hydrophyllic grasses, sedges and other aquatic plants
including some tree species like Terminalia arjuna,
Mitragyna parviflora etc., along their perimeters.
This ecosystem shows a zonation. The deeper ponding
area harbours "oating-leaved aquatic plants like
Nymphaea nouchali, Nelumbo nucifera, Nymphoides
spp., Aponogeton spp. in association with submerged
aquatics like Ceratophyllum demursum and Hydrilla
verticillata. Hygrorrhiza aristata, a wild relative of rice
forms dense buoyant mats while other "oating aquatics
like Pistia stratoites, Neptunia oleracea and Lemna
purpusilla are also found in inundated areas. Near the
edge of the villu, a range of grasses, forbs and sedges are
found. The characteristic species here are the succulent
grasses Iseilema laxum and Paspalidium flavidum.
The villus are important habitats for resident and
migrant water birds which include different kinds of
plovers, sandpipers, pipits, wag-tails, terns, teels, gulls,
geese, ducks, rails, eagles, harriers, cormorants, shags,
jacanas etc.
In the villus, both the vegetation and fauna have
been surveyed in the Mahaweli "ood plains110,111.
d) Seasonally flooded forests:
Seasonally "ooded marsh forests are known only
from one site, at Waturana, near Bulathsinhala112. This
ecosystem in the past would have been much more
widespread in the low-lying areas of the lowland wet
zone, but due to conversion of these habitats to paddy
cultivation, the Waturana site is possibly the last remnant
of this vegetation. The endemic rare and threatened plant
species recorded in this ecosystem include Stemonoporus
moonii, Mesua stylosa, Areca concina and Diospyros
quaesita; natural populations of the !rst two of these
species are now restricted to this site.
e) Streams, Rivers and Reservoirs:
Though small in size, Sri Lanka has 103 rivers (80
seasonal "owing through the dry zone, 22 perennial
Biodiversity of Sri Lanka 37
Journal of the National Science Foundation of Sri Lanka 36 Special Issue November 2008
Figure 11 : a) A villu rich in aquatic biota in the Mahaweli •ood plains near Dimbulagala rock outcrop in the background. b) There is a clear
zonation of the vegetation with •oating aquatic plants in the centre and sedges , shrubs and trees along the perimeter of the villu as seen
in this picture taken during the dry season.
Figure 12 : a) A rocky stream – Theligamu Oya near the North-eastern end and b) a waterfall at the South-eastern end of the Knuckles range.
Rapidly •owing fresh-water habitats of this nature harbour a biota unique to them.
(a) (b)
(a)
(b)
traversing the wet and intermediate zones, with the
325 km long Mahaweli alone running through all three
climatic zones) making up a collective length of 4,560
km (Figure 12).
These rivers, associated streams and waterfalls
provide special habitats for certain plant species in
the riverine and aquatic ecosystems, some like the
Cryptocoryne (all 10 species in Sri Lanka are endemic),
Laginandra (7 of the 8 species endemic) and Mapania
immersa are endemic to the island. Along the river or
stream banks, riverine or gallery forest formations can
be observed.
Studies carried out on the meso-fauna in mountain
streams at different elevations indicate a rich diversity of
species in areas of high currents113,114.
Sri Lanka does not have natural lakes, but around
12,000 reservoirs constructed for agricultural purposes
Nimal Gunatilleke et. al
38
Journal of the National Science Foundation of Sri Lanka 36 Special Issue
November 2008
during the past two millennia, ranging in size from
1–6500 ha, harbour a rich aquatic •ora and fauna115,116.
Sri Lanka’s species diversity: indigenous,
endemic and threatened species of fauna and
flora
In the island’s •ora, the highest species diversity is
recorded among the •owering plants, followed in
decreasing order by the fungi, bryophytes (mosses and
liverworts), freshwater algae and ferns (Table 2). Among
animals, the diversity of vertebrates is well known
compared to that of invertebrates, where only a few
groups have been studied in depth (Table 3).
Endemic species diversity: number and distribution
Plant Species: The endemic plant species diversity of
Sri Lanka comprises 927 or 28 % of •owering plants, of
which 60 % are found in the lowland wet zone and 34%
in the montane zone ecosystems of the island (Table 2).
Among the ferns, 59 species are endemic to Sri Lanka.
Of the island’s moss •ora, 11% are endemic 59,60. Recent
revisions on the island’s liverworts have not been
made and its endemic component is yet to be compiled.
Similarly, there is little information on the endemic
species among fungi, algae and lichens. In the lichen
family Thelotremataceae, as much as 32 % of the species
are endemic to the island and all of them are con!ned to
the rain forests125.
Animal Species: Among Sri Lanka’s ~ 930 vertebrates
species, 30 % are endemic to the island. The proportion of
endemic species among its amphibians is ~85 %, reptiles
~ 60 % and freshwater !shes 50 %. In the invertebrate
groups studied in depth, the endemic species component
among freshwater crabs is 100 %, land snails (83 %),
dragon•ies (47%) and butter•ies (8%) (Table 2). Like
the endemic plant species, most of the endemic fauna
too is con!ned to the wet-zone natural ecosystems of
Sri Lanka.
The increasing human-elephant con•ict in the island
is also an indication of the poor quality, fragmentation
and possibly the inadequate extents of suitable habitat to
support Sri Lanka’s largest threatened mammal species,
the elephant (Elephas maximus), in the protected areas
of Sri Lanka.
Threatened, point endemic and extinct
species of Sri Lanka
Plant Species: Currently, Sri Lanka has over 675
Flowering plants 3771(927) Mammals 91 (16) Bees 148 (21)
Gymnosperms 1 (0) Birds 482 (33) Ants 181 ( ??)
Ferns & fern allies 314 (59) Reptiles 171 (101) Butter•ies 243 ( 20)
Mosses 561 (63) Amphibians 106+ (90+) Spiders 501 ( ??)
Liverworts 227 (??) Fishes 82 (44) Land snails 246 (204)
Freshwater algae 560+ ( ?) (Fresh water only) Dragon •ies 120 ( 57)
Fungi 2260+ ( ?) Crabs* 51 ( 51)
Lichens 661 ( ?) Shrimps* 23 ( 07)
Indigenous plant
species
Total no.
of species
(endemics)
Vertebrate
species
Total no.
of species
(endemic)
Invertebrates
species
No. of species
(endemics)
* Fresh water only.
Sources: reference nos. 59,60,117-124.
Table 2. Species diversity among selected •ora and fauna and the number of endemics in each group
Flora
Flowering Plants 412(61) 252
Ferns 30 (53) 60
Vertebrates
Mammals 14 (88) 27
Birds 16 (48) 30
Reptiles 37 (37) 19
Amphibians 51 (57) 1
Fishes 20 (45) 8
Invertebrates
Land snails 32 (16) 1
Freshwater crabs 37 (73) 0
Dragon•ies 20 (35) 0
Butter•ies 13 (65) 53
Plant and animal
groups
No. and (%) of nationally threatened
species
Endemic Non-endemic
Table 3. Numbers of threatened species of selected groups of
plants and animals.
* Percentage calculated on the total number of endemics.
Source: reference no. 117.
Biodiversity of Sri Lanka 39
Journal of the National Science Foundation of Sri Lanka 36 Special Issue November 2008
Figure 13 : Rocky out-crops or inselbergs like the Kokagala in the dry zone, harbour plants and animals rarely encountered
in the plains below.
Figure 14 : The paddy fields in the valleys and home gardens often with a stratification analogous to that of the natural forest but substituted with
plants of traditional economic value in the lower slopes is a sustainable man-made ecosystem rich in agro-biodiversity in Sri Lanka. The
rain forests in the upper slopes were cleared during the period of European colonization in the 19th century for the cultivation of rubber
as seen in these photographs.
(a) (b)
Nimal Gunatilleke et. al
40
Journal of the National Science Foundation of Sri Lanka 36 Special Issue
November 2008
•owering threatened plant species among the 1099
assessed so far124 and 90 fern species assigned a
threatened status. Information on the threatened status of
species in other plant groups is lacking.
Animal species: Among animals 37 – 88 % of the
endemic species in the vertebrate groups and over 16
-73 % of species in the invertebrate groups have been
assessed as threatened117 (Table 3).
Sri Lanka’s has 108 point (or micro) endemic
species, each recorded only from a single site. They
include 5 mammal species, 4 agamid lizards, 15
amphibians, 3 freshwater !sh and 81 •owering plant
species (Appendix 1), almost all con!ned to the island’s
wet zone.
There are a large number of species, both plants
and animals, not collected during the past century117.
Many of these could be either extinct or on the brink of
extinction. Among the plants, over 30 fern species and
72, (including 42 endemics) of 1099 •owering plants
assessed are suspected to be extinct. Among animals
suspected extinct are 21 endemic amphibians 117 and
considered extinct in historical times are the gaur and
comb duck126, and up to 13 species of snakes. Among
wild relatives of agricultural crops, those of rice are
severely threatened due to habitat loss.
Genetic diversity
Information on genetic diversity of the natural •ora and
fauna of Sri Lanka is extremely limited. Studies carried
out on the genetic diversity of leopards indicate lower
genetic variation in the Sri Lankan populations compared
to those of Peninsular Malaysia127,128. Molecular
analyses have shown that the Sri Lankan leopard is a
distinct subspecies, one of ten in the world. Among the
Sri Lanka elephants relatively high levels of differentiation
in mtDNA were observed between the northern,
mid-latitude and southern regions of the island129.
Recent molecular analyses have shown that the
direct developing tree frogs of the genus Philautus
(Rhacophoridae) represents a large, endemic insular
radiation42. Molecular phylogenetic studies also
have shown that Sri Lankan caecilian fauna are of
monophyletic origin, those of the southernwestern
lowlands comprising a clade, possibly representing a
relatively recent divergence from those of the central
highlands130.
Among the plants, phylogenetic and population
genetic studies have been carried out on some members
of Dipterocarpaceae131-134 (Shorea and the endemic
genus Stemonoporus), Myrtaceae135 (Syzygium spp.) and
Annonaceae136 (Polyalthia spp. and Xylopia championi).
The proportion of the total genetic diversity among
populations of some of the Dipterocarp species resulting
from higher rates of out-crossing is shown to be similar
to the average reported in literature for tropical woody
species. Lower within-population genetic variation
for these species have been reported in logged-over
forest fragments, suggesting that forest degradation,
fragmentation and habitat size reduction have lead to
their genetic erosion131,132.
Bar reef 0 - 3 78.5 ≈100 mortality
(1993-1994)
Bar reef 7 - 8 - ≈100 mortality 14
Kandakuliya 0 – 5 22 Small colonies corals smothered
were present by Halimeda
Hikkaduwa Nature Res. 0 - 4 47.2 7 12
Hikkaduwa south of NR 7 - 13 Not estimated Not measured 17
Rumassala 1-5 45 19.6 Better recovery at
4-5 m depth; live
coral not estimated
Weligama 0 - 2 92 28 54
31
Several new colonies
noted; too sparse to
estimate percent
Table 4. Status of selected coral reefs in Sri Lanka before and after the El-Nino event of April-May 1998
Location Depth (m) Pre-bleaching
(% live coral)
1999 – 2000
(% live coral)
2001 – 2002
(% live coral)
Source: reference no. 142
Biodiversity of Sri Lanka 41
Journal of the National Science Foundation of Sri Lanka 36 Special Issue November 2008
Figure 15 : Sri Lanka is rich in micro-endemic species, such as the ones shown here, which are restricted largely to Morningside,
a small forest reserve east of the Sinharaja World Heritage Site. These species were all discovered serendipitously in the
course of exploration during the past 15 years, suggesting that significant hitherto unrecorded diversity may exist in
other little-explored parts of the island, such as the Peak Wilderness. a) Calotes desilvai; b) Ceratophora erdeleni;
c) Ceratophora karu; d) Crocidura hikmiya; e) Cyrtodactylus subsolanus; f) Perbrinckia rosae; g) Philautus
frankenbergi; h) Philautus poppiae (Photos courtesy of Madhava Meegaskumbura).
(a)
(c)
(e)
(g)
(b)
(d)
(f)
(h)
Nimal Gunatilleke et. al
42
Journal of the National Science Foundation of Sri Lanka 36 Special Issue
November 2008
The genetic diversity of wild relatives, traditional
cultivars and land races of crop species are being
estimated for both in-situ and ex-situ conservation in
both within and outside protected areas, in farmlands
and home gardens, gene banks, where appropriate, and
characterized primarily through the initiatives of the
Departments of Agriculture and Export Agriculture and
Plantation Crop Research Institutes137. The diversity
of crop genetic resources in different agro-ecological
regions of the island is quite high. About 645 species of
crop wild relatives of Sri Lanka have been catalogued in
the National Herbarium at Peradeniya.
Agricultural and medicinal species and their
genetic diversity
Agricultural plant diversity138 in the island includes Oryza
sativa (rice) with its 2,800 varieties/land races and !ve
wild relatives, 7 coarse-grain species and their traditional
cultivars of maize and sorghum; 14 grain legume species;
8 cucurbitaceous, 2 solonaceous and 4 other vegetable
(bean, okra, amaranth, chilli) species; 17 root and tuber
crop species. The economically useful spices are 8
species of Cinnamon, Elettaria cardamomum, 3 Piper
species (with 7 wild relatives), clove, nutmeg, betel nut,
vanilla, chilli, and ginger. Others of importance include
three major beverage species (tea, coffee and cocoa),
sugarcane, kitul plam, arecanut, tamarind, citronella, 3
species of oil crops and 2 !bre crops. The horticultural
species include banana with nine cultivars and two wild
relatives, citrus, and over 20 other fruit species. The
germplasm accessions of these wide range of crops are
kept in the seed gene bank and the !eld gene banks of
the Department of Agriculture and also ex situ, in village
home gardens137.
A total of 1414 species of Sri Lankan plants are
considered to be of value to the indigenous (ayurvedic)
medicine. Among them 50 are heavily used, 208
commonly used and 79 are threatened. A project has
been carried out to lay a foundation for conservation and
sustainable use of medicinal plants in Sri Lanka139.
Degradation of biodiversity
Sri Lanka and the Western Ghats of India have been
designated as one of the 34 global hotspots of the world1
based on the fact that both these regions together harbour
at least 1,500 species of vascular plants as endemics
and have lost at least 70% of their original habitat140. In
addition to habitat loss, most of the remaining habitats
of endemics are degraded. In this section, factors
responsible for degradation of ecosystem diversity are
examined.
Marine and maritime ecosystems: Degradation of
coastal ecosystems is due to natural processes (wave
action, sea level rise, tsunamis) and human activities
such as the mining of beach and river sand, inland coral
deposits and shore reefs; construction of groins, harbours,
jetties etc.; and improperly sited coastal buildings and
removal of coastal vegetation. Destruction of mangrove
ecosystems has been attributed to land reclamation,
urbanization, pond aquaculture, extraction of !rewood
and timber, in"ow of sediments and pollutants (sewage,
industrial ef"uents) due to poor land use management
in inland areas, desalination due to inland irrigation
projects; harvesting polychaetes for brood-stock feed;
drag-net operations during !shing and conversion to salt
pans4,141.
Surveys of coral reefs before, during and after the
El Niño event of April-May 1998, showed extensive
bleaching of corals, up to a depth of 8 m and drastic
reduction of associated !sh populations. Recovery has
been slow and variable (Table 4) due to overgrowth
of calcarious Halimeda sp. and !lamentous algal
species142,143.
The world’s !fth-largest quake in a century hit
South and Southeast Asia on the 26th of December 2004
unleashing a tsunami that affected Sri Lanka as well.
Habitats impacted included sand dunes, salt marshes,
mangroves and coastal woodlands and scrublands. The
vegetation has been impacted by the force of the wave,
inundation by saltwater and sand deposition. Most
vegetation types have shown recovery and regeneration
with time but how individual species in them have been
affected remains to be assessed144.
Terrestrial ecosystems: Man’s in"uence on natural
vegetation in the island, closely related to the long
history of land use in Sri Lanka, falls broadly into six
periods (prehistoric, proto-historic, dry zone civilization
(500BC–1200AD), late medieval: (1200AD–1500AD),
western colonial: (1505 AD –1947AD) and post
Independence: (since 1948AD). During each of these
periods different parts of the country have been the focus
of human activities, that have adversely affected their
natural vegetation145.
Natural forest ecosystems: The quality of the natural
forest ecosystems varies with the forest type. With the
exception of relatively small extents in inaccessible
areas, most lowland rain forests have been converted to
cash-crop agriculture145-147, (cinnamon, rubber, coconut
and tea) and in recent times selectively logged to supply
plywood and hardwood timber (Figure 14). Logging at
!rst was slow, using elephants and hand implements.
Biodiversity of Sri Lanka 43
Journal of the National Science Foundation of Sri Lanka 36 Special Issue November 2008
After the 1960s, mechanized logging using chainsaws,
skidders, timber jacks, and loaders have led to their
rapid degradation148 until logging of natural forest was
prohibite altogether by an administrative order in 1989.
Almost all the tropical dry mixed evergreen forests
are believed to be secondary in origin and about 500–800
years old (Figure 13). The distribution of numerous tanks
(small reservoirs) in Sri Lanka’s dry zone suggests that
dry zone forests were converted to shifting cultivation
and paddy lands at one time or another. Once the
reservoirs were abandoned, the forests returned with
time, accounting for their secondary origin27,149. These
forests have also been intensively and selectively logged
for their very valuable hardwood timber species such as
Ebony (Diospyros ebenum), Palu (Manilkara hexandra),
Satinwood (Chloroxylon swietenia), Pannakka
(Pleurostylia opposita) and they have also been exploited
for fuel wood23. In the last few decades, much of this
ecosystem has been converted to agriculture following
major irrigation schemes.
The lower montane and montane forests, due to
their inaccessibility, were less influenced by man during
the pre-colonial period. From 1833–1880 (during the
British period), about one-thirteenth (4,760 sq. km) of
the island’s land area, mostly the lower montane forests
were converted to coffee, cinchona and tea plantations147.
Timber from these clearings was used both internally
and for export charging a royalty. More recently, areas
close to the perimeter in the remaining forests have been
exploited for fuel wood and for urban and agricultural
expansion. Good pristine forests may still be seen in
parts of the Peak Wilderness Sanctuary, Hakgala Strict
Nature Reserve, Knuckles, Pidurutalagala and Horton
Plains. Mining for precious stones also has affected
some of these forests. Parts of the Central Highlands,
Knuckles range and Rakwana-Deniyaya range have been
planted with cardamom after clearing the understorey.
Although this practice has been curtailed in recent times,
the removal of undergrowth (including of the canopy
species), has led to the paucity of juveniles and saplings
for regeneration and restoration of the forest structure
and composition. The enigmatic die-back of patches of
montane forests, very evident in the Horton Plains and
Knuckles65,66 also lowers the quality of these forests,
jeopardizing the biodiversity they harbour (Figure 16).
The gaps created by burning and die-back are favourable
habitats for the native bracken (Pteridium revolutum);
in less exposed gaps the native Strobilanthes spp. and
non-native invasive species Aristea eckloni grow
abundantly.
It is also ironical that the relative proportion of
population increase in the island’s wet zone districts,
where nearly 90% of the endemic biodiversity is
harboured, is considerably greater compared to that in
the dry zone. The proportion of endemic flowering plants
in the wet zone is 5.71 per 100 sq km, a "gure 50 times
greater than that in the dry zone (0.11 per 100 sq km).
The human population in a comparable area in the wet
zone on the other hand is three times greater than that of
the dry zone, giving rise to severe human pressure on the
aseasonal wet zone forests.
There are numerous non-wood forest products
extracted from the natural forest ecosystems of
Sri Lanka23,150-155. Among them are all 10 species of
small and large diameter rattans (Calamus)154, which
includes 8 endemics, bamboo (Ochlandra stridula,
Bambusa bambos), many different species of medicinal
plants150 such as Bin Kohomba, Weniwelgeta, Kothala
Himbutu, Rasa Kinda (Munronia pinnata, Coscinium
fenestratum, Salacia reticulata and Tinospora cordifolia,
respectively); and food plants such as Kitul, Beraliya,
Hal, Wood Apple (Caryota urens, Shorea spp., Vateria
copallifera, and Limonia acidissima, respectively)154.
Other products for domestic use are also extracted, such
as roof thatching material from Beru (Agrostistachys
intramarginalis), resins from several different species
of Shorea, Hal and Kekuna (Canarium zeylanicum)153.
In species like cane or the medicinal vine veniwelgeta
(Coscinium fenestratum), removal of the whole plant
drastically reduces the adult population. Thereby fruits
are not available for regeneration of the species and
precipitating consequent decline.
The past half-century has seen a steady growth in
the collection of aquatic "sh and plants for the aquarium
trade, ornamental plants including orchids and ferns,
and wild relatives of agricultural and medicinal species.
Among fresh water "sh species, 75% of the indigenous
"sh, including 21 endemics, had been collected from the
wild. However, effective policing and enforcement of
export regulations have curtailed this to some extent. In
the inland aquatic habitats, the water plants collected
illicitly from streams include species of Cryptocoryne
(most of which are endemic) and Lagenandra. Arguably
the biggest threat, however, to aquatic animal life has
been the introduction (both deliberately for "sheries and
accidentally by hobbyists) of dozens of exotic species
of "sh and other aquatic organisms, which are now
established in the island’s waterways.
Sand removal
In almost all of Sri Lanka’s major rivers sand is mined at
an alarmingly high rate for commercial purposes despite
augmented regulations to curtail this lucrative activity.
Nimal Gunatilleke et. al
44
Journal of the National Science Foundation of Sri Lanka 36 Special Issue
November 2008
Ecosystem Plant species Animal species
Thorn Scrub: Bundala National Park (NP) Prosopis juliflora, Feral buffalo
Opuntia dillenii
Savannas: Victoria-Randenigala-Rantambe sanctuary, Lantana camara Feral buffalo
Udawalawe National Park.
Stream & river banks: Lowland wet zone Dillenia suffruticosa, Golden apple snail
Anona glabra
Intermediate zones Mimosa pigra
Lower montane forests Clusia rosea, Wedelia trilobata
Miconia calvescens
Moist semi-evergreen forests Bambusa bambos
Lowland wet zone rain forests Wedelia trilobata,
Clidemia hirta,
Clerodendrum quadriloculare
(potentially invasive)
Montane Zone
Wet Patana Grasslands (Horton Plains) Ulex europaeus Common crow
Montane forests (Horton Plains) Cestrum aurantiacum,
Ageratina riparia,
Aristea eckloni
Marsh and/or aquatic ecosystems Salvinia molesta, Golden apple snail,
Eichhornia crassipes Thilapia, Carp
Table 5: Some invasive species in different natural ecosystems in Sri Lanka
Figure 16: Forest die-back in the a) Knuckles region near the Corbett's Gap area and the undergrowth invaded by a native bamboo species
b) Horton Plains National Park.
(b)
(a)
Biodiversity of Sri Lanka 45
Journal of the National Science Foundation of Sri Lanka 36 Special Issue November 2008
This not only damages the river bed and pollutes the
water, but greatly reduces the amount of sand being
carried to the seashore. Consequently, the sea sand
washed away due to wave action is not compensated for
by the in•ow of sand brought in by the rivers leading to
severe coastal erosion.
Competition by invasive exotic and indigenous
species
Over the years, a large number of exotic plant and
animal species have been introduced, either accidentally
or purposefully into Sri Lanka. Some of these species
have escaped into natural ecosystems and are now not
only growing successfully, but also out-competing the
indigenous species in them (Table 5).
In a few other instances some indigenous species
in their natural habitats have rapidly expanded their
populations, changing their vegetation structure, species
composition and even reducing their species richness.
Such examples include the bamboo species, Davidsea
attenuata (Bata) in parts of the Knuckles range, where
it is fast replacing the typical montane tree species
and Bambusa bambos (Katu Una) in the Victoria-
Randenigala-Rantambe sanctuary and in forests near
Bakamuna, out-competing the characteristic species of
these tropical moist semi-evergreen forests. Population
explosions of both invasive and even indigenous species
are indicators of the quality and health of the ecosystems
and a good understanding of the long-term dynamics
of these systems is imperative if these changes are to be
reversed.
Lack of understanding of scientific management
Scienti!c management of almost all natural ecosystems in
the island at present is still in its inception. Until recently,
management of the forest and grassland ecosystems had
been largely limited to protection, commercial extraction
of timber and control of visitor activities. In most of the
inland aquatic habitats, management has been completely
ignored. Monitoring the good health of these ecosystems
and taking timely measures to counteract any adverse
changes have hardly received any attention. This, among
other reasons is now posing many problems to Sri
Lanka’s natural ecosystems. A few may be highlighted
here:
In Horton Plains, the sambar (Cervus unicolor)
population has increased, possibly in response to the more
preferred fodder156-157 (Pennisetum clandestinum Kikiyu
grass, a naturalized exotic, and P. glabrum), which has
been inadvertently introduced during the period when
part of the area was cultivated for seed potato production.
Especially during lactating periods (July-September and
November-January) of sambar, the grazing pressure is
higher on the carpet grasses, richer in nitrogen than on the
bushy tussock grass Chryzopogon nodulibarbis. This, if
unchecked may lead to over-browsing of preferred tree
species (Cinnamomum ovalifolium, Neolitsea fuscata,
Calophyllum walkeri, Eurya japonica, Elaeocarpus
subvillosus, Syzygium revolutum and Hedyotis trimenii)
in the natural forest.
Continuing forest die-back and the spread of
invasive species appear to alter the physiognomy and
species composition of the forest landscape. Increasing
number of vehicles (hence emissions) and visitors to the
reserve entail problems of garbage disposal resulting
in the increasing the abundance of the common crow,
washing vehicles and discharging pollutants into streams.
Predation of the indigenous fauna, by crows, also calls
for attention.
Air pollutants, agrochemicals and vehicle exhaust
too can adversely affect biodiversity. Lichens are good
indicators of air pollution. Their diversity and abundance
greatly decrease with pollution, and those species that
withstand pollution may increase their populations, out-
competing those that are sensitive to pollution.
Acid rain has also been reported in Horton Plains158.
The extent to which it contributes to forest die-back
and water quality in this ecosystem needs to be
investigated.
In Bundala, the increasing population of feral
buffalos competing with other herbivores for fodder
and freshwater, rapid invasion of the thorn scrub by
Prosopis juliflora and Opuntia dillenii at the expense
of indigenous species, and changing salinity levels in
lagoons due to the Lunugamvehera irrigation project,
need to be addressed urgently.
In the Udawalawe National Park and Victoria-
Randenigala-Rantembe Sanctuary control of !re has
resulted in the invasion of the savannas by Lantana
camara followed by other tree species. This ground-
layer woody vegetation is less attractive to herbivores
compared to the grass cover which it had replaced.
In the arid zone, as in Yala, the long dry season is a
natural tool to restrain the scrub, unless aggressive
weedy species like P. julifera and O. dilenii take over.
Graminivorous birds and other fauna and •ora dependent
on the grassland habitat would also be affected adversely
if timely remedial measures are not taken to maintain
ecosystem and habitat diversity in these reserves.
Nimal Gunatilleke et. al
46
Journal of the National Science Foundation of Sri Lanka 36 Special Issue
November 2008
Pollution
With the increasing conversion of land to agriculture in
Sri Lanka, indiscriminate application of agrochemicals
is widespread. Due to the relatively high rainfall, some
of these agrochemicals eventually pollute waterways
and reservoirs. Industrial waste disposal into waterways
also contributes to the poor quality of the island’s
water resources. Thereby, the richness of the aquatic
fauna and •ora may be impacted. On many occasions
algal blooms and •sh kills have been reported in the
Kothmale Reservoir and more recently in the Kandy
Lake. Indirectly through the effect of pesticides on food
chains, the natural biodiversity of these systems may be
reduced.
Sediment loads in the water also affect the
biodiversity of aquatic systems. Sediment not only
blocks out the light, adversely affecting photosynthesis
by primary producers in the system, but by reducing
visibility in the water impede their foraging and mating
activities.
In protected areas where visitor pressure is high, solid
waste, particularly polythene waste, is often a problem.
Animals have been seen feeding at garbage dumps and
consuming polythene in or near protected areas some
times with fatal consequences.
Over visitation
During the last two decades the number of visitors,
including school children to wildlife reserves and natural
ecosystems have increased many-fold, compounding
the problems of solid waste, vehicular traf•c and noise
pollution, all of which unless carefully managed could
lead to degradation of these ecosystems.
Introduction of motorized boats to the Madu Ganga
and other aquatic systems have had many adverse effects.
Among them are bank erosion due to wave action and oil
pollution. The extensive use of kerosene powered vessels
and boats using two-stroke engines in the Hikkaduwa
Marine Sanctuary has been identi•ed as a major threat
to the reef.
Table 6. Important policies, international conventions and legislation related to biodiversity conservation in Sri Lanka.
A. National Policies
i. National Forestry Policy (1995)
ii. National Wildlife Conservation Policy (1990)
B. International BD related Conventions signed by Sri Lanka
i. Biological Diversity (CBD)
ii. International Trade in Endangered Species of wild •ora and fauna (CITES)
iii. Protection of the World Cultural and Natural Heritage
iv. Wetlands of international importance (Ramsar)
v. Conservation of Migratory Species (Bonn Convention)
vi. Control of Persistent Organic Pollutants,
vii. Combating Deserti•cation,
viii. UN Framework Convention on Climate Change,
C. Laws & Regulations Directly/Indirectly Related to BD Conservation
i. Fauna and Flora Protection Ordinance 2 (1937) and Amendment Acts 44 (1964) 1 (1970), 49 (1993) and 2 (1996).
ii. Forest Ordinance No. 3 (1945), Amendment Acts No. 13 (1966) and 13 (1988)
iii. National Wilderness Heritage Act No. 3 (1988)
iv. Fisheries and Aquatic Resources Act No. 2 (1996)
v. National Environment Act 47 (1980) and Amendment Act 56 (1988)
vi. Crown Land Ordinance (1929)
vii. National Aquatic Resources Research and Development Agency Act No.54 (1981) and Amendment Act 32 (1996).
viii. The Land Development Act No. 10 (1983) and Amendment Acts 22 (1993), 9 (1995), 20 (1996).
ix. Soil Conservation Act (1951)
x. Coast Conservation Act No. 57 (1981) and Amendment Act No. 64 (1988).
xi. Botanic Gardens Ordinance 3 (1961)
xii. National Zoological Gardens Act 41 (1982)
xiii. Plant Protection Act No. 35 (1999)
xiv. Animal Diseases Act. 59 (1992)
xv. Aired Act 31 (1996) and Amendment Acts No. 5 (1962) and 9 (1969), 7 (1977), 6 (1978).
Biodiversity of Sri Lanka 47
Journal of the National Science Foundation of Sri Lanka 36 Special Issue November 2008
Grasslands
While parts of the wet patana grasslands have been
converted to agriculture (Horton Plains, Nuwara Eliya,
Knuckles), some dry patana grasslands in the Uva basin
are gradually becoming urbanized, or giving way to,
agriculture and exotic mono-speci!c forest plantations.
Many thalawas, damanas and villus have been subject
to grazing, and some converted to cash-crop agriculture.
These ecosystems have also been subject to !re, a factor
essential to maintain them. But recurrent !res at short
intervals lead to increased erosion, soil impoverishment
and soil moisture stress, all leading to colonization by
exotic invasive weeds that aggressively compete with the
native species.
Legal and institutional framework for
conversation
Biodiversity conservation policy in Sri Lanka is based
on the National Wildlife Policy (2000) and the National
Forestry Policy (1995). Several other policy statements
too, impinge on the biodiversity sector, including
the National Environment Policy and the National
Ecotourism Policy. The legal framework for biodiversity
conservation rests primarily with the Fauna and Flora
Protection Ordinance (FFPO) of 1937 with its later
revisions, the Forest Ordinance, National Wilderness
Heritage Act, National Environmental Act, Fisheries
Ordinance, Coast Conservation Act and several other
legislative acts. While some of these acts are overlapping
in their mandate, others are, to some extent, in con"ict as
seen in the case of FFPO and the Fisheries and Aquatic
Resources Act in respect of the import and export of !sh
and other aquatic organisms159. The FFPO employs two
approaches for species conservation: i) establishment
and maintenance of a protected area system wherein all
biota within the speci!ed areas are protected from human
interference, and ii) listing of protected species in seven
different schedules indicating taxonomic groups that are
protected (positive listing) or not protected (negative
listing)160.
Sri Lanka is also a party to several international
legal instruments/conventions relating to biodiversity
conservation (Table 6). For the export of wild biota, a
permit is required unless otherwise stated in regulations.
Sri Lanka is bound by the Convention on International
Trade in Endangered Species of Wild Fauna and Flora
(CITES) which seeks international co-operation in
regulating the growing threat from the trade in endangered
biota.
Based on the government policies, complying
with appropriate international legal instruments and
national regulations on biodiversity which are revised
from time to time, a number of sectoral master plans,
action plans and strategies have been prepared and are
being implemented by the relevant government agencies
(primarily the Departments of Wildlife Conservation
and Forest Conservation) in collaboration with civil
society.
The document titled ‘Biodiversity Conservation
in Sri Lanka: A Framework for Action’159 prepared by
the Ministry of Forestry and Environment in responding
to the article 6 of the Convention on Biological
Diversity outlines the national strategies proposed
for the conservation, sustainable management and
equitable transfer of bene!ts arising from the biological
wealth and its associated knowledge in Sri Lanka. In
addition, sector based environmental action programmes
including those on forestry, wildlife conservation,
agriculture, plantations, !sheries, coastal and marine
area management and tourism for the period 2003-2007
have been spelt out161.
Biodiversity conservation: some future
directions
From the above review of literature on the biodiversity of
Sri Lanka, it is clearly evident that this island is blessed
with a rich and diverse biota distributed in a wide array
of landscapes and waterscapes. In addition, Sri Lanka
also possesses an equally rich religio-cultural heritage
spanning over the millennia which had instilled a strong
conservation ethic amongst its people. Nonetheless,
the degradation of biodiversity has been quite severe
particularly during the last two centuries and this trend
is still continuing in many different ways in spite of
having a comprehensive array of enactments based on
sound policies for the conservation of biodiversity. This
suggests that mere enactment of regulations without their
strict enforcement had not been an effective measure
for the conservation of biodiversity and its sustainable
management. The regulations to be enforced effectively,
there need a strong motivation and commitment
towards conservation among all stakeholders which
can come effectively through increased awareness of
the conservation value through !eld based conservation
education and awareness programmes. The policy and
legal framework also fails to recognize that protective
measures alone cannot save species threatened with
extinction. Despite over 100 species being listed as
threatened, there has up to now been very little effort to
devise and implement recovery plans for any of these.
At the Millennium Summit of the United Nations
held in New York in 2000, eight Millennium Development
Nimal Gunatilleke et. al
48
Journal of the National Science Foundation of Sri Lanka 36 Special Issue
November 2008
Goals (MDGs) were targeted for achieving during the
decade from 2005-2015. Among these, there are at least
two viz., achieving universal primary education (goal #2)
and ensuring environmental sustainability (goal #7) that
could be used effectively in biodiversity conservation in
the longer term. The Convention on Biological Diversity
(CBD) also aims at achieving its Biodiversity Target by
reducing the rate of biodiversity loss by 2010. Taking
these global proclamations further at the World Summit
on Sustainable Development held in Johannesburg
in 2002, the governments agreed to reorient national
education systems to a vision of sustainability that links
economic well-being with respect to cultural diversity
and the resources of the planet.
Responding to these challenges, the UNESCO
launched its interdisciplinary approaches to help
governments and policy makers to achieve these goals
through actions aimed at:
i. Sustaining biodiversity at UNESCO World Heritage
Sites and Biosphere Reserves,
ii. Linking traditional knowledge to biodiversity
management and governance,
iii. Building partnerships to prevent biodiversity loss
and degradation, and
iv. Building dialogue to prevent conflicts and share
bene"ts.
The World Network of Biosphere Reserves is developed
as a mean to integrate human development with
biodiversity conservation and sustainable management
of natural resources. As such, they serve as learning
platforms and ‘out-door laboratories’ for testing and
demonstrating innovative approaches to reconcile
conservation of biodiversity with social and economic
development through enhancing cooperation among
academic, political, practitioner and other stakeholder
communities. The United Nations Decade for
Sustainable Development (DESD) for the period from
2005-2014 emphasizes the critical role of education in
achieving sustainable development 162. The 3rd World
Congress of Biosphere Reserves held in February
2008 at which the Madrid Action Plan (MAP) for the
period 2008-2012 was presented, has clearly identi"ed
the need for research, training, capacity building and
demonstration agendas on conservation and sustainable
use of biodiversity, mitigation and adaptation to climate
change and socio-economic and cultural well-being of
human communities.
In Sri Lanka too, the network of biosphere
reserves (both national and international) established
in representative ecosystems could be effectively used
for meeting the challenges described above. From the
primary education levels, appreciation of nature is being
instilled in children and the secondary school curriculum
now encourages all students to carry out investigative
projects. For students with interest in biodiversity
conservation, the out-door laboratories of the country’s
ecosystems are a valuable resource yet to be tapped in
a meaningful and innovative manner in meeting our
national obligations to the above international goals.
Education centers, nature trails and nature interpretation
programmes at these sites need to be developed to
realize the full potential of these ecosystems as out-door
laboratories.
The protected areas network and their surrounding
landscape with human habitation in Sri Lanka could
be managed in accordance with the MAB concept
which intends to ful"l three complementary functions:
conservation of biological and cultural diversity,
sustainable development and logistic support for
education, training, research, and monitoring. To ful"l
these functions, the biosphere reserves are organized in
three interrelated zones: a legally constituted core area
devoted to long-term protection, a buffer zone where
activities compatible with the conservation objectives can
take place, and an outer transition area where sustainable
resource management practices are promoted and
developed 163. This would provide a pragmatic approach
for nature conservation in its fullest sense in Sri Lanka
as practiced in pre-colonial times while implementing
the national conservation strategies and also meeting
national obligations to international conventions on
biodiversity conservation and environment protection.
Biodiversity of Sri Lanka 49
Journal of the National Science Foundation of Sri Lanka 36 Special Issue November 2008
Appendix 1: ‘Point endemic’ species, each known only from a single site < 100 km2 (for terrestrial sites) or
< 5 linear km (for aquatic sites, streams and rivers). Plant records based on Dassanayake & Fossberg, 1980–2004.
Reptiles other than agamids are excluded for lack of data recorded from Sri Lanka.
Group/Family/Species Locality Group/Family/Species Locality
BIRDS None FLOWERING PLANTS
AGAMID LIZARDS Acanthaceae
Barleria nutans Hantana
Gymnostachyum thwaitesii Ambagamuwa
Strobilanthes arnottiana Nuwara Eliya
Strobilanthes caudata Hantana
Strobilanthes hypericoides Gartmore
Strobilanthes nigrescens Ambagamuwa
Strobilanthes punctata Peak Wilderness
Strobilanthes thwaitesii Ambagamuwa
Anacardiaceae
Semecarpus moonii Kottawa
Annonaceae
Gonicthalamus thomsonii Haycock
Apocynaceae
Wrightia flavido-rosea Doluwakanda
Aquifoliaceae
Ilex knucklesensis Knuckles
Araceae
Arisaema constrictum Ramboda
Cryptocoryne alba Ingiriya
Asclepiadaceae
Brachystelma lankana Laggala
Burmanniaceae
Thismia gardneriana Kuruwita
Celastraceae
Maytenus fruticosa Nawalapitiya
Clusiaceae
Mesua stylosa Kalutara
Compositae
Anaphalis fruiticosa Peak Wilderness
Gynura hispida Horton Plains
Cornaceae
Mastixia congylos Knuckles
Mastixia montana Hakgala
Mastixia nimalii Sinharaja
Cyperaceae
Eleocharis lankana Yakkala
Mapania immersa Sinharaja
Scleria pilosa Kalutara
Dipterocarpaceae
Hopea brevipetiolaris Doluwakanda
Stemonoporus reticulatus Kanneliya
Agamidae
Cnemaspis podihuna Lahugala
Ceratophora tennentii Knuckles
Ceratophora karu Morningside
Ceratophora erdeleni Morningside
AMPHIBIANS
Bufonidae
Adenomus dasi Peak Wilderness
Ranidae
Limnonectes greenii Horton Plains
Microhylidae
Microhyla karunaratnei Morningside
Rhacophoridae
Philautus macropus Knuckles
Philautus limbus Haycock
Philautus alto Horton Plains
Philautus simba Morningside
Philautus fulvus Knuckles
Philautus caeruleus Peak Wilderness
Philautus lunatus Handapanella Pl.
Philautus papillosus Handapanella Pl.
Philautus procax Morningside
Philautus ocularis Morningside
Philautus nemus Haycock
FRESHWATER FISH
Cyprinidae
Puntius bandula Galapitamada
Puntius asoka Kitulgala
Puntius srilankensis Pallegama
Nimal Gunatilleke et. al
50
Journal of the National Science Foundation of Sri Lanka 36 Special Issue
November 2008
Appendix 1. FLOWERING PLANTS continued
Group/Family/Species Locality Group/Family/Species Locality
Ebenaceae Orchidaceae continued
Diospyros albiflora Morapitiya Oberonia fornicata Knuckles
Diospyros atrata Gannoruwa Oberonia wallie-silvae Rangala
Diospyros koenigii Gannoruwa Oberonia weragamaensis Weragama
Diospyros oblongifolia Kanneliya Robiquetia virescens Knuckles
Diospyros oppositifolia Haycock Taeniophyllum gilimalense Gilimale
Ericaulaceae Orobanchaceae
Eriocaulon subglaucum Horton Plains Christisonia albida Hakgala
Euphorbiaceae Legocia aurantiaca Hakgala
Glochidion gardneri Peradeniya Piperaceae
Phyllanthus dealbatus Rassagala Peperomia species Bopathalawa
Phyllanthus hakgalensis Hakgala Poaceae
Phyllanthus oreophilus Horton Plains Arundinaria densifolia Horton Plains
Gentianaceae Arundinaria scandens Pidurutalagala
Crawfurdia championii Horton Plains Arundinella thwaitesii Ambagamuwa
Gesneriaceae Garnotia fuscata Sabaragamuwa
Didymocarpus floccosus Dolosbage Garnotia panicoides Culloden Estate
Loranthaceae Heteropholis nigrescens Hakgala
Dendrophthoe lonchiphyllus Ambagamuwa Ischaemum polystachyum Matale
Helixanthera ensifolia Karawita Kanda Oplismenus thwaitesii Nalanda
Macrosolen barlowii Horton Plains Zenkeria obtusiflora —
Melastomataceae Podostemaceae
Memecylon gracillimum Doluwakanda Farmeria metzgerioides Hakkinda
Memecylon orbiculare Kalubowitiyakanda Rhamnaceae
Mimosaceae Ziziphus lucida Erankeliya Eraniya
Acacia lankaensis Diyaluma Rosaceae
Albizia lankaensis Frotoft Estate Sanguisorba indicum Peak Wildern.
Monimiaceae Rutaceae
Hortonia ovalifolia Peak Wilderness Zanthoxylum caudatum Eratne
Myrsinaceae Scrophulariaceae
Ardisia wightiana Athweltota Adenosma subrepens Hewesse
Myrtaceae Umbelliferae
Eugenia glabra Naunkitaela Peucedanum ceylanicum Belihul Oya
Orchidaceae Urticaceae
Dendrobium maccarthiae Kuruwita Elatostema walkerae Ramboda
Gastrodia zeylanica Haycock Zingiberaceae
Oberonia claviloba Ambagamuwa Amomum acuminatum Ekneligoda
Oberonia dolabrata Knuckles Amomum benthamianum Reigam Korale
Amomum graminifolium Sinharaja
Amomum nemorale Hewesse
Biodiversity of Sri Lanka 51
Journal of the National Science Foundation of Sri Lanka 36 Special Issue November 2008
FLOWERING PLANTS
Acanthaceae
Acanthus illicifolius L.
Barleria nutans Nees
Gymnostachyum thwaitesii T. Anders.
Strobilanthes arnottiana Nees
Strobilanthes caudata T. Anders.
Strobilanthes hypericoides J. R. I. Wood
Strobilanthes nigrescens T. Anders.
Strobilanthes punctata Nees
Strobilanthes thwaitesii T. Anders.
Anacardiaceae
Nothopegia beddomei Gamble
Mangifera zeylanica (Blume) Hook. F.
Semecarpus moonii Thw.
Annonaceae
Anona glabra L.
Goniothalamus thomsonii Thw.
Xylopia championii Hook. f. & Thoms.
Apocynaceae
Wrightia flavido-rosea Trimen
Aquifoliaceae
Ilex knucklesensis Philcox
Arecaceae
Areca concinna Thw.
Caryota urens L.
Nypa fruticans Wurmb
Araceae
Arisaema constrictum Barnes
Cryptocoryne alba de Wit
Pistia stratiotes L.
Asclepediaceae
Calotropis gigantea (L.) R. Br.
Asclepiadaceae
Brachystelma lankana Dassanayake & Jayasuriya
Asteraceae
Ageratina riparia (Regel) King & H. Rob.
Anaphalis fruiticosa Hook. f.
Elephantopus scaber L.
Gynura hispida Thw.
Wedelia trilobata (L.) A.S. Hitchc.
Avicenaceae
Avicennia marina (Forssk.) Vierh.
Avicennia officinalis L.
Bombacaceae
Cullenia rosayroana Kosterm.
Cullenia zeylanica (Gardner)K. Schum.
Burmanniaceae
Thismia gardneriana Hook. f. ex Thw.
Burseraceae
Canarium zeylanicum (Retz.) Blume
Cactaceae
Opuntia dillenii (Ker-Gawl.) Haw.
Celastraceae
Maytenus fruticosa (Thw.) Loes.
Pleurostylia opposita (Wall.) Alston
Hippocrateaceae
Salacia reticulata Wight
Chenopodiaceae
Halosarcia indica (Willd.) P. G. Wilson
Salicornia brachiata Roxb.
Sueda maritima (L.) Dumort.
Sueda monoica Forssk. Ex J. F. Gmelin
Clusiaceae
Calophyllum inophyllum L.
Calophyllum trapezifolium Thw.
Calophyllum walkeri Wight
Clusia rosea Jacq.
Garcinia hermonii Kosterm.
Mesua ferrea L.
Mesua stylosa (Thw.) Kosterm.
Combretaceae
Lumnitzera racemosa Willd.
Terminalia arjuna (Roxb.) Wight &Arn.
Terminalia bellirica (Gaertn.) Roxb.
Terminalia chebula Retz.
Convolvulaceae
Evolvulus alsinoides (L.) L.
Ipomea pes-caprae (L.) R. Br.
Cornaceae
Mastixia congylos Kosterm.
Mastixia montana Kosterm.
Mastixia nimalii Kosterm.
Cyperaceae
Eleocharis lankana T. Koyama
Mapania immersa (Thw.) Benth. ex Clarke
Scleria pilosa Boeckeler
Appendix 2 : Authority names of plant and animal species referred to in the paper, including tables and Appendix 1.
Nimal Gunatilleke et. al
52
Journal of the National Science Foundation of Sri Lanka 36 Special Issue
November 2008
Datiscaceae
Tetrameles nudiflora R. Br.
Dilleniaceae
Dillenia suffruticosa (Griffith) Martelli
Dipterocapaceae
Dipterocarpus hispidus Thw.
Dipterocarpus zeylanicus Thw.
Hopea brevipetiolaris (Thw.) Alston
Vateria copallifera (Retz.) Alston
Stemonoporus moonii Thw.
Shorea gardneri (Thw.) Ashton
Shorea trapezifolia (Thw.) Ashton
Stemonoporus reticulatus Thw.
Ebenaceae
Diospyros melanoxylon Roxb.
Diospyros albiflora Alston
Diospyros atrata Alston
Diospyros ebenum Koenig.
Diospyros koenigii Kosterm.
Diospyros oblongifolia (Thw.) Kosterm.
Diospyros oppositifolia Thw.
Diospyros quaesita Thw.
Elaeocapaceae
Elaeocarpus subvillosus Arn.
Ericaceae
Rhododendron arboreum ssp. zeylanicum (Booth)
Tagg
Ericaulaceae
Eriocaulon subglaucum Ruhland
Euphorbiaceae
Agrostistachys intramarginalis Philcox
Dimorphocalyx glabellus Thw.
Drypetes sepiaria (Wight & Arn.) Pax & Hoffm.
Glochidion gardneri Thw.
Phyllanthus dealbatus Alston
Euphorbiaceae Contd.
Phyllanthus emblica L.
Phyllanthus hakgalensis Thw. ex Trimen
Phyllanthus oreophilus Muell. Arg.
Phyllanthus polyphyllus Willd.
Fabaceae
Acacia lankaensis Kosterm.
Acacia leucophloea (Roxb.) Willd.
Acacia planifrons Wight & Arn.
Albizia lankaensis Kosterm.
Alisicarpus vaginalis (L.) DC.
Bauhinia racemosa Lam.
Butea monosperma (Lam.) Taub.
Cassia auriculata L.
Desmodium heterophyllum (Willd.) DC.
Desmodium triflorum (L.) DC.
Dichrostachys cinerea (L.) Wight & Arn.
Mimosa pigra L.
Neptunia oleracea Lour.
Pericopsis mooniana (Thw.) Thw.
Prosopis juliflora (Sw.)DC.
Pterocarpus marsupium Roxb.
Ulex europaeus L.
Gentianaceae
Crawfurdia championii (Gardner)Trimen
Exacum walkeri Arn.
Gesneriaceae
Didymocarpus floccosus Thw.
Hydrocharitaceae
Enhalus acoroides (L. f.) Royle
Hydrilla verticillata (l. f.) Royle
Thalassia hemprichii (Ehrenb.) Asch.
Iridaceae
Aristea eckloni Baker
Lauraceae
Actinodaphne speciosa Nees
Cinnamomum ovalifolium Wight
Cryptocarya wightiana Thw.
Litsea ovalifolia (Wight) Trimen
Neolitsea fuscata (Thw.) Alston
Lecythidaceae
Barringtonia asiatica (L.) Kurz
Careya arborea Roxb.
Lemnaceae
Lemna purpusilla Torrey
Loranthaceae
Dendrophthoe lonchiphyllus (Thw.) Danser.
Helixanthera ensifolia (Thw.) Danser.
Macrosolen barlowii Wiens
Malvaceae
Thespesia populnea (L.) Sol.ex Correa
Melastomataceae
Clidemia hirta (L.) D. Don
Memecylon gracillimum Alston
Melastomataceae
Memecylon orbiculare Thw.
Miconia calvescens DC.
Osbeckia octandra (L.) DC.
Meliaceae
Munronia pinnata (Wall.) Theob.
Biodiversity of Sri Lanka 53
Journal of the National Science Foundation of Sri Lanka 36 Special Issue November 2008
Menispermaceae
Coscinium fenestratum (Gaertn.) Colebr.
Tinospora cordifolia (Willd.) Hook. f. & Thoms.
Monimiaceae
Hortonia ovalifolia Wight
Moraceae
Antiaris toxicaria Leschen.
Artocarpus gomezianus Wall. Ex Trecul
Myristicaceae
Myristica dactyloides Gaertn.
Myrsinaceae
Ardisia wightiana (Wall. Ex A. DC.) Mez.
Myrtaceae
Eugenia bracteata Roxb.
Eugenia glabra Alston
Psidium guajava L.
Syzygium revolutum Walp.
Syzygium rotundifolium Arn.
Syzygium umbrosum Thw.
Nelumbonaceae
Nelumbo nucifera Gaertn.
Nymphaeaceae
Nymphaea nouchali Burm. f.
Orchidaceae
Dendrobium maccarthiae Thw.
Gastrodia zeylanica Schlechter
Oberonia claviloba Jayaweera
Oberonia dolabrata Jayaweera
Oberonia fornicata Jayaweera
Oberonia wallie-silvae Jayaweera
Oberonia weragamaensis Jayaweera
Robiquetia virescens (Gard. ex Lindley) Jayaweera
Satyrium nepalense D. Don
Taeniophyllum gilimalense Jayaweera
Orobanchaceae
Christisonia albida Thw. ex Benth.
Legocia aurantiaca (Wight) Livera
Poaceae
Themeda triandra Forssk.
Andropogon lividus Thw.
Bamboosa bambos (L.) Voss ex Vilmorin
Sinarundinaria densifolia (Munro). Chao & Renvoize
Aristida setacea Retz.
Arundinaria scandens Soderstrom & Ellis
Arundinella thwaitesii Hook. f.
Arundinella villosa Arn. ex Steud.
Chrysopogon aciculatus (Retz.) Trin.
Chrysopogon nodulibarbis (Steud.) Henrard
Cyanodon dactylon (L.) Pers.
Cymbopogon nardus (L.) Rendle
Cymbopogon polyneuros (Steud.) Stapf.
Davidsea attenuata(Thw.) Soderstrom & Ellis
Echinocloa colona (L.) Link
Garnotia fuscata Thw.
Garnotia panicoides Trimen
Garnotia exaristata Gould
Heteropholis nigrescens (Thw.) C. E. Hub.
Hygrorrhiza aristata (Retz.) Nees.
Imperata cylindrica (L.) Rausch.
Ischaemum polystachyumJ. & C. Presl.
Iseilema laxum Hack.
Ochlandra stridula Moon ex Thw.
Oplismenus thwaitesii Hook. f.
Oryza sativa L.
Panicum maximum Jacq.
Paspalidium flavidum (Retz.) A. Camus
Pennisetum clandestinum Hochst. Ex Chivo.
Pennisetum glabrum Steud.
Spinifex littoreus (Burm. f. ) Merr.
Sporobolus diander (Retz.) P. Beauv.
Themeda tremula (Steud.) Hack.
Zenkeria obtusiflora (Thw.) Benth.
Podostemaceae
Farmeria metzgerioides (Trimen) Willis ex.
Hook. f.
Pontideriaceae
Eichhornia crassipes (Mart.) Solms-Laub.
Rhamnaceae
Ziziphus oenoplia (L.) Miller
Ziziphus lucida Moon ex Thw.
Rhizophoraceae
Bruguiera gymnorhiza (L.) Savigny
Bruguiera sexangula (Lour.) Poir.
Ceriops tagel (Perr.) C. B. Rob.
Rhizophora apiculata Blume
Rhizophora mucronata Poir.
Rosaceae
Sanguisorba indicum (Gardner) Tirv.
Rubiaceae
Mitragyna parviflora (Roxb.) Korth.
Hedyotis trimenii Deb & Dutta
Rutaceae
Chloroxylon swietenia D.C.
Limonia acidissima L.
Zanthoxylum caudatum Alston
Salvadoraceae
Salvadora persica L.
Sapindaceae
Dimocarpus longan Lour.
Felicium decipiens (Wight & Arn.) Thw.
Schleichera oleosa (Lour.) Oken
Nimal Gunatilleke et. al
54
Journal of the National Science Foundation of Sri Lanka 36 Special Issue
November 2008
Sapotaceae
Manilkara hexandra (Roxb.) Daubard
Scrophulariaceae
Adenosma subrepens (Thw.) Benth.
Pedicularis zeylanica Benth.
Solonaceae
Cestrum aurantiacum Lindley
Sonneratiaceae
Sonneratia alba J. Sm.
Sonneratia caseolaris (L.) Engl.
Sterculiaceae
Pterospermum suberifolium (L.) Willd.
Pterygota thwaitesii (Masters) Alston
Symplocaceae
Symplocos cochinchinensis (Lour.) S. Moore
Theaceae
Eurya chinensis R. Br.
Thymelaeaceae
Wikstroemia indica (L.) C. A. Meyer
Ulmaceae
Gironniera parvifolia Planch.
Apiaceae
Peucedanum ceylanicum Gardner
Urticaceae
Elatostema walkerae Hook. f.
Vacciniaceae
Gaultheria leschenaultii DC.
Verbenaceae
Clerodendrum quadriloculare (Blanco) Merr.
Lantana camara L.
Zingiberaceae
Amomum acuminatum Thw.
Amomum benthamianum Trimen
Amomum graminifolium Thw.
Amomum nemorale (Thw.) Trimen
Elettaria cardamomum (L.) Maton
PTERIDOPHYTES
Ceratophyllaceae
Ceratophyllum demursum L.
Dennstaedtiaceae
Pteridium revolutum (Blume) Nakai
Salviniaceae
Salvinia molesta D.S. Mitch.
FAUNA
MAMMALS
Elephantidae
Elephas maximus Linnaeus, 1758
AGAMID LIZARDS
Agamidae
Cophotis ceylanica Peters, 1861
Calotes nigrilabris Peters, 1860
Cnemaspis podihuna Deraniyagala, 1944
Ceratophora tennentii Gunther & Gray 1861
Ceratophora karu Pethiyagoda & Manamendra-
Arachchi, 1998
Ceratophora erdeleni Pethiyagoda & Manamendra-
Arachchi, 1998
SEA TURTLES
Cheloniidae
Caretta caretta (Linnaeus, 1758)
Chelonia mydas (Linnaeus, 1758)
Eretmochelys imbricata (Linnaeus, 1766)
Lepidochelys olivacea (Eschscholtz, 1829)
Dermochelyidae
Dermochelys coriacea (Vandelli, 1761)
AMPHIBIANS
Bufonidae
Adenomus dasi Manamendra-Arachchi & Pethiyagoda,
1998
Ranidae
Fejervaryia greenii (Boulenger, 1904)
Limnonectes greenii (Boulenger, 1904)
Microhylidae
Microhyla karunaratnei Fernando & Siriwardhane, 1996
Microhyla zeylanica Parker & Hill, 1949
Rhacophoridae
Philautus alto Manamendra-Arachchi & Pethiyagoda,
2005
Philautus caeruleus Manamendra-Arachchi &
Pethiyagoda, 2005
Philautus fulvus Manamendra-Arachchi & Pethiyagoda,
2005
Philautus limbus Manamendra-Arachchi & Pethiyagoda,
2005
Philautus lunatus Manamendra-Arachchi &
Pethiyagoda, 2005
Philautus macropus (Gunther, 1869)
Philautus microtympanum (Gunther, 1859)
Philautus nemus Manamendra-Arachchi & Pethiyagoda,
2005
Philautus ocularis Manamendra-Arachchi &
Pethiyagoda, 2005
Philautus papillosus Manamendra-Arachchi &
Biodiversity of Sri Lanka 55
Journal of the National Science Foundation of Sri Lanka 36 Special Issue November 2008
Pethiyagoda, 2005
Philautus poppiae Meegaskumbura & Manamendra-
Arachchi, 2005
Philautus procax Manamendra-Arachchi & Pethiyagoda,
2005
Philautus schmarda (Kelaart, 1854)
Philautus simba Manamendra-Arachchi & Pethiyagoda,
2005
Philautus steineri Meegaskumbura & Manamendra-
Arachchi, 2005
Philautus variabilis (Gunther, 1859)
Polypedates eques Gunther, 1858
FRESH WATER FISHES
Cyprinidae
Puntius bandula Kottelat & Pethiyagoda, 1991
Puntius asoka Kottelat & Pethiyagoda, 1989
Puntius srilankensis (Senanayake 1985)
FRESH WATER SHRIMPS AND CRABS
Atyidae
Lancaris kumariae (de Silva, 1990)
Lancaris sinhalensis (Ortmann, 1894)
Parathelphusidae
Ceylonthelphusa sorror (Zehntner, 1894)
Ceylonthelphusa diva Bahir & Ng, 2005
Ceylonthelphusa savitriae Bahir & Ng, 2005
Perbrinckia glabra Ng, 1995
Perbrinckia gabadagei Bahir & Ng, 2005
Perbrinckia punctata Ng, 1995
Perbrinckia rosae Bahir & Ng, 2005
ANTS
Aneuretinae
Aneuretus simoni (Emery, 1893)
References
1. Mittermeier R.A., Gil P.R., Hoffmann M., Pilgrim
J., Brooks T., Goettsch Mittermeier C., Lamoreux J. &
Da Fonseca G.A.B. (2004). Hotspots Revisited: Earth’s
Biologically Richest and Most Endangered Terrestrial
Ecoregions. pp. 392. Cemex, Mexico.
2. Kotagama S.W. & Bambaradeniya C.N.B. (2006). An
Overview of the Wetlands of Sri Lanka and Their
Conservation Significance. pp. 11. IUCN Sri Lanka and
the Central Environmental Authority, Colombo,
3. Anonymous (2002). State of the Environment in
Sri Lanka. A National Report Prepared for the South Asian
Association for Regional Cooperation. pp. 245. Ministry
of Environment and Natural Resources, Battaramulla.
4. Anonymous (1990). Coastal Zone Management Plan. pp.
81. Coast Conservation Department, Colombo.
5. Abeywickrama B. A. & Arulgnanum. P. (1993). The
Coastal Plants of Sri Lanka Part I (Salt Marsh Plants) pp.
19. National Science Council of Sri Lanka, Colombo 7.
6. Pemadasa M. A., Balasubramaniam S., Wijewansa H. G.
& Amarasinghe L. (1979). The ecology of a salt marsh in
Sri Lanka. Journal of Ecology 67(1): 41 - 63.
7. Arulchelvam K. (1968). Mangroves. The Ceylon Forester
8 (3 & 4): 52-92.
8. Ranawana K. B. (1994). Ecology of lagoon fringing and
riverine mangroves of the Northwest, West and South
Coasts of Sri Lanka. M. Phil. Thesis, University of
Peradeniya, Peradeniya.
9. Bambaradeniya C. N. B., Ekanayake S. P., Kekulandela
L. D. C. B., Samarawickrama V. A. P., Ratnayake N. D. &
Fernando R. H. S. S. (2002). An assessment of the Status
of Biodiversity in the Muthurajawela Wetland Sanctuary.
pp. 48. Occassional Papers of IUCN Sri Lanka, No. 3.
10. Jayatissa L.P., Dahdouh-Guebas F. & Koedam N. (2002).
A review of the floristic composition and distribution of
mangroves in Sri Lanka. Botanical Journal of the Linnean
Society 138: 29 - 43.
11. Amerasinghe M. D. (1989). Structural and functional
properties of mangrove ecosystems in Puttalam Lagoon
and Dutch Bay, Sri Lanka. M. Phil. Thesis, University of
Peradeniya, Peradeniya.
12. De Silva K. H. G. M. & Balasubramaniam, S. (1984/1985).
Some ecological aspects of the mangrroves on the west
coast of Sri Lanka. Ceylon Journal of Science (Biological
Sciences). 17/18: 22-41.
13. Pinto L. (1978). Some ecological principles of selected
mangrove islets in Negombo Lagoon (Sri Lanka).
M. Phil. Thesis, University of Colombo, Colombo 03.
14. Kugathassan K. S. (1969). Mangrove vegetation of the
lagoon. A guide to the study of distribution, identi"cation
and characterization of the lagoon in particular and
common species, in general. Hydrobiological survey
of the Thondamannaru Lagoon. Bulletin No. 7 pp. 23.
Northern Province Teachers Association, Sri Lanka.
15. de Silva K. H. G. M. & de Silva P. K. (1998). Status,
diversity and conservation of the mangrove forests
of Sri Lanka. Journal of South Asian Natural History
3 (1): 79 -102.
Nimal Gunatilleke et. al
56
Journal of the National Science Foundation of Sri Lanka 36 Special Issue
November 2008
Organization of the United Nations.
31. Gunatilleke C.V.S., Gunatilleke I.A.U.N., Ethugala
A. U. K. & Esufali S. (2004). Ecology of Sinharaja Rain
Forest and the Forest Dynamics Plot in Sri Lanka’s
Natural World Heritage Site. pp. 221, Wildlife Heritage
Trust Publications (Pvt.) Ltd., Colombo.
32. Wijesinghe M.R. (2006). Ecological traits of endemic
small mammals in rainforests of Sri Lanka and their
implications for conservation. In: Fauna of Sri Lanka:
Status of Taxonomy Research and Conservation. (Ed.
C.N.B. Bambaradeniya) pp. 232-234, The World
Conservation Union, Colombo and Government of Sri
Lanka.
33. Kotagama S.W. & Goodale E. (2004). The composition
and spatial organisation of mixed-species flocks in a
Sri Lankan rainforest. Forktail 20: 63-70.
34. Goodale E. & Kotagama S.W. (2005). Testing the roles
of species in mixed-species bird flocks of a Sri Lankan
rainforest. Journal of Tropical Ecology 21: 669-676.
35. Goodale E. & Kotagama S.W. (2007). Some observations
on the geographic variation of mixed-species bird flocks
in Sri Lanka. Journal of the Bombay Natural History
Society 104: 96-98.
36. De Silva A. (2006). Current status of the reptiles of
Sri Lanka. In: Fauna of Sri Lanka: Status of Taxonomy
Research and Conservation. (Ed. C.N.B. Bambaradeniya)
pp. 134-163, The World Conservation Union, Colombo
and Government of Sri Lanka.
37. Pethiyagoda R. (1994). Threats to the indigenous
freshwater "shes of Sri Lanka and remarks on their
conservation. Hydrobiologia 285:189-201.
38. Wikramanayake E.D. & Moyle P.B. (1989). Ecological
structure of tropical "sh assemblages in wet zone streams
of Sri Lanka. Journal of Zoology, London 218: 503-526.
39. Amerasinghe U.S., Shirantha R.R.A.R. & Wijeratne
M. J. S. (2006). Some aspects of ecology of endemic
freshwater "shes of Sri Lanka. In: Fauna of Sri Lanka:
Status of Taxonomy Research and Conservation
(Ed.C.N.B. Bambaradeniya) pp. 113 – 124, The World
Conservation Union, Colombo and Government of Sri
Lanka.
40. Pethiyagoda R. & Manamendra-Arachchi K. (1998).
Evaluating Sri Lanka’s amphibian diversity. Occasional
Papers of the Wildlife Heritage Trust 2: 1-12.
41. Manamendra-Arachchi K. & Pethiyagoda R. (2005). The
Sri Lankan shrub frogs of the genus Philautus Gistel, 1848
(Ranidae: Rhacophorinae), with description of 27 new
species. In: Contributions to Biodiversity Exploration in
Research in Sri Lanka. (Eds. D.C.J. Yeo, P.K.L. Ng & R.
Pethiyagoda) The Raffles Bulletin of Zoology, Supplement
No. 12: 5 - 145.
42. Meegaskumbura M., Bossuyt F., Pethiyagoda R.,
Manamendraarachchi K., Bahir M., Milinkovitch M.C. &
Schneider C.J. (2002). Sri Lanka: an amphibian hotspot.
Science 298 (5592): 379
43. Perera W. P. N. & Bambaradeniya C. N. B. (2006).
Species richness, distribution and conservation status of
butterflies in Sri Lanka. In: Fauna of Sri Lanka: Status
of Taxonomy Research and Conservation. (Ed. C.N.B.
16. Pinto L. (1980). Some ecological aspects of a population
of mangrove crabs occurring within the islets of Negombo
Lagoon (Sri Lanka). Proceedings of the Asian Symposium
on Mangrove Environment, Research and Management,
August 25-29, Kuala Lumpur.
17. Pinto L. & Wignarajah S. (1980). Some ecological aspects
of the edible oyster Crassostrea cuculata (Born) occuring
in association with the mangroves in the Negombo
Lagoon, Sri Lanka. Hydrobiologia 69: 11 - 20.
18. Pinto L. & Punchihewa N. N. (2004). Utilisation of
mangroves and seagrasses by "shes in the Negombo
Estuary, Sri Lanka. Marine Biology 126(2): 333 - 345.
19. Samarakoon J. & van Zon H. (1991). Environmental
Profile of Muthurajawela and Negombo Lagoon. Greater
Colombo Economic Commission and Euroconsult, The
Netherlands.
20. Amerasinghe U. S., Amerasinghe M. D. & Nissanka C.
(2002). Investigation of the Negombo Estuary (Sri Lanka)
brush park "shery, with an emphasis on community-based
management. Fisheries Management and Ecology 9 (1):
41-56.
21. De Silva A. (2006). An annotated bibliography of
publications on marine turtles of Sri Lanka. Indian Ocean
Turtle Newsletter 3: 12-26.
22. Ashton P.S. & Gunatilleke C.V.S. (1987). New light on the
plant geography of Ceylon I: Historical plant geography.
Journal of Biogeography 14: 249-285.
23. Anonymous (1995). Sri Lanka Forestry Sector Master
Plan. pp. 511. Forestry Planning Unit, Ministry of
Agriculture, Battaramulla.
24. Gunatilleke I.A.U.N. & Gunatilleke C.V.S. (1984).
Distribution of endemics in the tree flora of a lowland
hill forest in Sri Lanka. Biological Conservation 28:
275–285.
25. Gunatilleke C.V.S. & Ashton P.S. (1987). New light
on the plant geography of Ceylon II: The ecological
biogeography of the lowland endemic tree flora. Journal
of Biogeography 14: 295-327.
26. Gunatilleke C.V.S. & Gunatilleke I.A.U.N. (1985).
Phytosociology of Sinharaja: a contribution to rain forest
conservation in Sri Lanka. Biological Conservation
31:21-40.
27. Peeris C.V.S. (1975). The ecology of the endemic tree
species of Sri Lanka in relation to their conservation.
Ph.D. Thesis, University of Aberdeen, UK.
28. Singhakumara B.M.P. (1995). Ecological Assessment
of the Kanneliya-Dediyagala-Nakiyadeniya (KDN)
Forest Complex. pp. 70, Department of Forestry
and Environmental Science, University of Sri
Jayewardenepura, Gangodawila, Nugegoda.
29. Manage U. (1999). Structure and floristic survey of the
Delwala Proposed Reserve with emphasis on altitudinal
variation. B.Sc. Research Report, University of Colombo,
Colombo 03.
30. Anonymous (1997). Designing an Optimum Protected
Areas System for Sri Lanka’s Natural Forests, Vol. 1,
pp. 201. The World Conservation Union, World
Conservation Monitoring Centre and Food and Agriculture
Biodiversity of Sri Lanka 57
Journal of the National Science Foundation of Sri Lanka 36 Special Issue November 2008
Bambaradeniya) pp.53-64. The World Conservation
Union, Colombo, and Government of Sri Lanka.
44. d’ Abrera B. (1998). The Butterflies of Ceylon. pp 224.
Wildlife Heritage Trust, Sri Lanka, 95, Cotta Road,
Colombo 8.
44a. Gamage R. (2007). An Illustrated Guide to the Butterflies
of Sri Lanka. pp 264. Tharanjee Prints, Nawinna.
45. Benjamin S. P. & Bambaradeniya C. N. B. (2006).
Systematics and conservation of spiders in Sri Lanka:
current status and future prospects. In: Fauna of Sri Lanka:
Status of Taxonomy Research and Conservation. The
World Conservation Union, Colombo and Government of
Sri Lanka.
46. Dias R.K.S. (2002). Current knowledge on ants of
Sri Lanka. ANet Newsletter 4 : 17-20.
47. Gunawardene N.R., Daniels A. E. D., Gunatilleke I. A.
U. N., Gunatilleke C. V. S., Karunakaran P. V., Nayak
K. G., Prasad S., Puyravaud P., Ramesh B. R., Subramanian
K. A. & Vasanthy G. (2007). A brief overview of the
Western Ghats - Sri Lanka biodiversity hotspot. Current
Science 93(11): 1567 - 1572.
48. Bahir M.M., Ng P.K.L., Crandall K. & Pethiyagoda R.
(2005). A conservation assessment of the freshwater
crabs of Sri Lanka. In: Contributions to Biodiversity
Exploration and Research in Sri Lanka. (Eds. D.C.J. Yeo,
P.K.L. Ng & R. Pethiyagoda). The Raffles Bulletin of
Zoology, Supplement No.12: 121 – 126.
49. Naggs F.D., Ranawana K. & Mapatuna Y. (2005). The
Darwin Initiative Project on Sri Lankan land snails:
patterns of diversity in Sri Lankan forests.Contributions
to Biodiversity Exploration in Research in Sri Lanka.
(Eds. D.C.J. Yeo, P.K.L. Ng & R. Pethiyagoda) The
Raffles Bulletin of Zoology, Supplement No.12: 23 – 29.
50. Naggs F.D. & Raheem D. (2000). Land Snail Diversity in
Sri Lanka. Natural History Museum, London. CD ROM.
51. Greller A.M., Gunatilleke I.A.U.N., Jayasuriya A.H.M.,
Gunatilleke C.V.S., Balasubramaniam S. & Dassanayake,
M.D. (1987). Stemonoporus dominated montane forests
in the Adam’s Peak Wilderness, Sri Lanka. Journal of
Tropical Ecology 3(1):243-253.
52. Jayasuriya A.H.M., Greller A.M., Balasubramaniam
S., Gunatilleke C.V.S., Gunatilleke I.A.U.N. &
Dassanayake M.D. (1993). Phytosociological studies of
the mid-elevational (lower montane) evergreen forests
in Sri Lanka. In: Ecology and Landscape Management
in Sri Lanka pp. 79-94 (Eds. W. Erdelen, C. Preu, N.
Ishwaran & C.M. Madduma Bandara) Proceedings of the
International and Interdisciplinary Symposium, Colombo,
Weikersheim:Margraf, Germany.
53. Sinhakumara B.M.P. (1995). Floristic Survey of Adam’s
Peak Wilderness. pp 156. Forest Department of Sri Lanka,
Battaramulla.
54. Gunatilleke I.A.U.N., Greller A.M., Jayasuriya
A.H.M., Gunatilleke C.V.S. & Balasubramaniam
S. (1996). Vegetation of the Peak Wilderness and its
conservation. Phyta (Journal of the Botanical Society,
University of Peradeniya) 4(1): 1-9.
55. Gunatilleke I.A.U.N., Gunatilleke C.V.S. & Dilhan
M.A.A.B. (2005). Plant biogeography and conservation
of the South-western hill forests of Sri Lanka. The Raffles
Bulletin of Zoology, Supplement No.12: 147-160.
56. Kostermans A. J. G. H. (1992). A Handbook of the
Dipterocarpaceae of Sri Lanka. pp. 224. Wildlife Heritage
Trust of Sri Lanka, 95, Cotta Road, Colombo 08.
57. Rubasinghe S.C.K. (2007). Species limits and
phylogenetics of the endemic genus Stemonoporus
(Dipterocapaceae). M.Phil. Thesis, University of
Peradeniya, Peradeniya.
58. Ekanayake S.P. (1994). A Phytosociological study of
semi-evergreen forests of the Knuckles and Udawalawe,
Sri Lanka. M.Phil. Thesis, University of Peradeniya,
Peradeniya.
59. O’Shea B.J. (2002). Checklist of the mosses of Sri Lanka.
Journal of the Hattori Botanical Laboratories 92: 125 –
164.
60. O’Shea B. J. (2003). Bryogeographical relationships of
the mosses of Sri Lanka. Journal of the Hattori Botanical
Laboratories 93: 293 – 304.
61. Balasubramaniam S., Ratnayake S. A. & White
R. (1993). The montane forests of the Horton
Plains Nature Reserve. In: Ecology and Landscape
Management in Sri Lanka (Eds. W. Erdelen, C. Preu,
N.Ishwaran, C.M. Madduma Bandara) pp. 95-108.
Proceedings of the International and Interdisciplinary
Symposium, Colombo, Weikersheim:Margraf, Germany.
62. Wijesundara D.S.A. (1991). Phytosociology of a montane
forest in Sri Lanka. M. Phil Thesis, University of
Peradeniya, Peradeniya.
63. Ratnayake H.D. (2005). Some aspects of the vegetation
in the Knuckles ecosystem. Lyriocephalus Special Issue
6(1 & 2):233 - 266.
64. Perera W.R.H. (1978). Thotapolakanda: an environmental
disaster. The Sri Lanka Forester 13 (3 &4): 53-55.
65. Werner W. L. (1988). Canopy die-back in the upper
montane rain forests of Sri Lanka. Geojournal 17 (2):
245-248.
66. Adikaram N.K.B., Ranawana K.B. & Weerasuriya
A. (2006). Forest Die Back in the Horton Plains National
Park. pp. 54. Department of Wildlife Conservation, No.
18, Gregorys Road, Colombo 07.
67. Werner W.L. (1982).The upper montane forests of
Sri Lanka. The Sri Lanka Forester 15: 119 - 135.
68. de Silva A., Ranawana K. B. & Goonewardene S. (2005).
Some observations of the mollusks of the Knuckles
massif. In: The Diversity of the Dumbara Mountains.
(Ed. A. de Silva) Lyriocephalus Special Issue 6 (1 &2).
179 - 182.
69. de Silva A. (2006). Current status of the reptiles of Sri
Lanka. In: Fauna of Sri Lanka: Status of Taxonomy
Research and Conservation. (Ed. C.N.B. Bambaradeniya)
pp. 134-163. The World Conservation Union, Colombo,
and Government of Sri lanka.
70. Manamendra-Arachchi K. & Pethiyagoda R. (2005).
The Sri Lankan shrub frogs of the genus Philautus Gistel,
1848 (Ranidae: Rhacophorinae), with description of 27
Nimal Gunatilleke et. al
58
Journal of the National Science Foundation of Sri Lanka 36 Special Issue
November 2008
new species. The Raffles Bulletin of Zoology, Supplement
No. 12: 163 - 303.
71. Meegaskumbura M. & Manamendra-Arachchi K . (2005).
Description of eight new species of shrub frogs (Ranidae:
Rhacophorinae: Philautus) from Sri Lanka. The Raffles
Bulletin of Zoology, Supplement No. 12: 305 - 338.
72. Cai Y. & Bahir M. M. (2005). Lancaris, a new genus
of freshwater shrimp from Sri Lanka (Crustaceae:
Deacapoda: Atyidae). The Raffles Bulletin of Zoology,
Supplement No. 12: 39-46.
73. Bahir M. & Ng P.K.L. (2005). Description of ten
new species of freshwater crabs (Crustacea:
Brachyura: Parathelphusidae: Ceylonthelphusa, Mahatha,
Perbrinckia) from Sri Lanka. The Raffles Bulletin of
Zoology, Supplement 12: 47-75.
74. Sriyani W. M., Padmalal U. K. G. K. & Kotagama
S. (2005). A study of mixed species bird •ocks of
submontane wet evergreen forest in the northern •ank
of the Knuckles region in Sri Lanka. In: The Diversity
of the Dumbara Mountains. (Ed. A. Silva) Lyriocephalus
Special Issue 6 (1 &2): 277 - 284.
75. Nizam B.Z., Padmalal U.K.G.K. & Kotagama S. (2005).
Amphibian species diversity along the northern •ank
of the Knuckles range Sri Lanka. In: The Diversity of
the Dumbara Mountains. (Ed. A. Silva) Lyriocephalus
Special Issue 6(1&2): 267 – 273
76. de Silva A. (2005). The Diversity of the Dumbara
Mountains. Lyriocephalus Special Issue 6 (1 &2): 382.
77. Ashton P.S. (1980). Dipterocarpaceae. In: A
Revised Handbook to the Flora of Ceylon. Vol.1.
(Eds. M. D. Dasanayake & F. R. Fosberg), pp. 364 -423.
78. Ranawana K.B., Ekanayake S.P., Gunatilleke I.A.U.N.,
Gunatilleke C.V.S., Herath H.M.D.R., Jayasena
H. A. H., Perera W., Rajapakse H. A. & Ranasingha
M. (1997). Resource Inventories for Protected areas:
Victoria - Randenigala- Rantambe Sanctuary. A Global
Environment Facility Project. Final Report submitted
to the Department of Wildlife Conservation, Sri Lanka.
pp. 310
79. Gunawarddena N.P.T. & Gunatilleke C.V.S. (1996).
The understorey vegetation at Samanalawewa in Sri
Lanka. Phyta (Journal of the Botanical Society, University
of Peradeniya) 4 (1): 18-33.
80. Weerasinghe, U.R. (1999). Seasonal resource use of large
herbivorous mammals in the Wasgomuwa National Park,
Sri Lanka. Ph. D. Thesis, University of Tokyo, Japan.
81. Jayasingam T. & Vivekanantharaja S. (1994). Vegetation
survey of the Wasgomuwa National Park, Sri Lanka:
analysis of the Wasgomuwa Oya Forest. Plant Ecology
113 (1): 1-8.
82.
Jayasingam T., Vivekanantharajah K. & Balasubramaniam
S. (1992). Vegetation survey of the Wasgomuwa National
Park: Reconnaissance. Vegetatio 101: 171-181.
83. Liyanage S. (1998). Floristic structure and aspects of
regeneration of tree species in the Kahalla Forest Reserve.
M. Phil. Thesis, University of Peradeniya, Peradeniya.
84. Fernando H.S.K. (1996). A comparative study on the
ecology of woody vegetation of forest types in the
Maduru Oya National Park. M. Phil Thesis, University of
Peradeniya, Peradeniya.
85. Jayasuriya A.H.M. (1980). Studies on the •ora and
ecology of Ritigala Strict Natural Reserve. M. Phil.
Thesis, University of Peradeniya, Peradeniya.
86. Jayasuriya A.H.M. (1984). Flora of Ritigala Natural
Reserve. Sri Lanka Forester 16: 60-156.
87. Jayasuriya A.H.M. & Pemadasa M.A. (1983). Factors
affecting the distribution of tree species in a dry zone
montane forest in Sri Lanka. Journal of Ecology 71:
571-583.
88. Dilhan M.A.A.B. (2005). Vegetation structure and
•oristic composition in the irrigation extension area of
the Lower Walawe Basin, Sri Lanka. M.Phil. Thesis,
University of Peradeniya, Peradeniya.
89. Dittus W.P.J. (1985). The in•uence of cyclones on the dry
evergreen forest of Sri Lanka. Biotropica 17: 1-14.
90. Dittus W.P.J. (1977). The ecology of a semi-evergreen
forest community in Sri Lanka. Biotropica 9: 268-286.
91. Dilhan M.A.A.B., Yakandawela D., Gunatilleke C.V.S.,
& Bambaradeniya C.N.B. (2002). Structure and
composition of scrubland vegetation in the Lower
Walawe Basin irrigation extension area in Sri Lanka.
Ceylon Journal of Science 30: 105 -124.
92. Sathurusinghe A. (1985). Phytosociological analysis of
vegetation of Wilpattu National Park. M. Sc. Dissertation
University of Sri Jayawardenapura, Gangodawila,
Nugegoda.
93. de Silva M. & de Silva P.K. (2004). The Yala Wildlife
Reserves: Biodiversity and Ecology. pp. 227, Wildlife
Trust Publications Ltd., Colombo.
94. Mueller-Dombois D. (1972). Crown distortion and
elephant distribution in the woody vegetation of Ruhuna
National Park Ceylon. Ecology 53(2): 208-226.
95. Bambaradeniya C.N.B., Ekanayake S.P., Fernando
R.H.S.S., Perera W.P.N. & Somaweera R. (2002). A
biodiversity status pro!le of Bundala National Park:
a Ramsar Wetland in Sri Lanka. Occasional Papers of
IUCN Sri Lanka No.2, pp. 37.
96. Mueller-Dombois D. (1968). Ecogeographic analysis
of a climate map of Ceylon with special reference to
vegetation. Ceylon Forester 8: 39 - 58.
97. Pemadasa M. A. (1984). Grasslands. In: Ecology and
Biogeography in Sri Lanka. (Ed. C.H. Fernando) pp. 453-
492. W. Junk Publishers.
98. de Silva A. (2007). The Diversity of Horton Plains
National Park (with Special Reference to its
Herpetofauna). pp. 273. Vijitha Yapa Publications,
Colombo.
99. Mueller-Dombois D. & Perera Malini. (1971). Ecological
differentiation and soil fungal distribution in the montane
grasslands of Ceylon. Ceylon Journal of Science
9(1): 1-41.
100. Pemadasa M. A. & Mueller Dombois D. ( 1981). An
association-analysis of montane grasslands of Sri Lanka.
Australian Journal of Ecology 6: 111-121.
101. Amarasinghe L. (1979).Ecology of grassland vegetation
of Hantana hills. M. Phil. Thesis, University of Peradeniya,
Peradeniya.
Biodiversity of Sri Lanka 59
Journal of the National Science Foundation of Sri Lanka 36 Special Issue November 2008
102. Pemadasa M.A. & Amarasinghe L. (1982). Ecology of a
montane grassland in Sri Lanka I. Quantitative description
of the vegetation. Journal of Ecology 70: 1-15.
103. Ashton P.M., Samarasinghe S.J., Gunatilleke I.A.U.N. &
Gunatilleke C.V.S. (1997). Role of legumes in release of
successionally arrested grasslands in the Central Hills of
Sri Lanka. Restoration Ecology 5(1): 36-43.
104. Kularatne H.A.G.C., Herath H.M.P.C., Gunatilleke
C.V.S. & Gunatilleke I.A.U.N. (1998). Towards park
management of the Victoria-Randenigala-Rantembe
(VRR) Sanctuary: Post-!re early successional vegetation
dynamics. Proceedings of the Final Workshop of the
University of Peradeniya-Oxford Forestry Institute Link
Project. pp. 127-152.
105. Kularatne H.A.G.C., Gunatilleke I.A.U.N. & Gunatilleke
C.V.S. (1996). Phenology of some savanna tree species at
Victoria-Randenigala Sanctuary. (Ed. H.P.M. Gunasena).
Proceedings of the Phase I Review sSeminar Workshop
of the UP-OFI Link Project. University of Peradeniya,
pp. 3-32.
106. Kularatne H.A.G.C., Gunatilleke C.V.S., Gunatilleke
I.A.U.N. & Brown N.D. (1998). Towards park
management of the Victoria-Randenigala-Rantembe
(VRR) Sanctuary: soil seed bank studies in the !re
affected sites. (Ed. H.P.M. Gunasena). Proceedings of the
Final Workshop of the University of Peradeniya-Oxford
Forestry Institute Link Project. pp. 102-126.
107. Russel-Smith J., Karunaratne N. & Mahindapala R. (2006).
Rapid inventory of wild medicinal plant populations in
Sri Lanka. Biological Conservation 132(1): 22-32.
108. Szechowyez R.W. (1961 ). The savanna forest of the
Gal Oya catchment (Ceylon). The Ceylon Forester
5(1&2): 17-22.
109. Koelmeyer K.O. (1957). Climatic classi!cation and the
distribution of vegetation in Ceylon. The Ceylon Forester
3(2): 144-163.
110. Jansen M. (1981). Villus of the "oodplains of the
Mahaweli ganga. Loris 15(6): 337-340.
111. Jansen M. (1987). Wetland habitats of the Mahaweli
ganga "ood plain. Loris 17(5): 186-191.
112. Seneviratne L. K., Ginige P., Dayananda M. D. &
Gamlath D. (1994). Wetland Site Report, Walauwatta
Waturana Swamp Forest Wetland Conservation Project.
pp. 54. Central Environmental Authority (Sri Lanka)/
Euroconsult, The Netherlands.
113. Starmulhner F. (1984). Mountain stream fauna with
special reference to the mollusca. In. Ecology and
Biogeography of Sri Lanka. (Ed. C.H. Fernando)
pp. 215-252.
114. Costa C.H. (1984). The ecology and distribution of
free living meso and macrocrustacea of inland waters.
In: Ecology and Biogeography of Sri Lanka. (Ed. C.H.
Fernando) pp. 195-213.
115. Schiemer F., Amarasinghe U.S., Frouzova J.,
Sricharoendham B. & Silva E.I.L. (2001). Ecosystem
structure and dynamics – A management basis for
Asian reservoirs and lakes. In: Reservoir and culture-
based !sheries: biology and management. Proceedings
of an International Workshop, February 15-18, 2000,
Bangkok. held in Bangkok, ACIAR Proceedings
No. 98: 215-226.
116. Fernando C.H. & de Silva S.S. (1984). Man made
lakes: ancient heritage and modern biological resource.
In: Ecology and Biogeography of Sri Lanka. (Ed. C.H.
Fernando). pp. 431-451.
117. The World Conservation Union, Sri Lanka (2007). The
2007 List of Threatened Fauna and Flora of Sri Lanka.
pp. 148. IUCN, 53, Horton Place, Colombo 07.
118. Shafer-Fehre M. (2006). A Revised Handbook to the
Flora of Ceylon. Vol. 15 (Part A:) Ferns and Fern-Allies.
pp.310. Science Publishers, En!eld, USA.
119. Abeywickrama B.A. (1979). The Genera of the Fresh
Water Algae of Sri Lanka Part 1. pp. 103. National
Science Council of Sri Lanka, 47/5, Maitland Place,
Colombo 07.
120. Coomaraswamy U. (1979a). A Handbook to the Fungi
Parasitic on the Plants of Sri Lanka pp. 169. National
Science Council of Sri Lanka. 47/5, Maitland Place,
Colombo 07.
121. Coomaraswamy U. (1979b). A Handbook to the Agaricus
of Sri Lanka, pp 68. National Science Council of Sri
Lanka, 47/5, Maitland Place, Colombo 07.
122. Coomaraswamy U. (1983). A Handbook to the Fungi
Associated with Insects of Sri Lanka, pp. 72. National
Science Council of Sri Lanka, 47/5, Maitland Place,
Colombo 07.
123. Coomaraswamy U. & Kumarasingham S. (1988). A
Handbook to the Macrofungi of Sri Lanka, pp. 124.
Natural Resources, Energy and Science Authority, Sri
Lanka, 47/5, Maitland Place, Colombo 07.
124. Bambaradeniya C.N.B. (2006). Fauna of Sri Lanka:
Status of Taxonomy, Research and Conservation. pp.
308. The World Conservation Union, Colombo and
Government of Sri Lanka.
125. Hale M. E. (1981). A revision of the lichen family
Thelotremataceae in Sri Lanka. Bulletin of the
British Museum of Natural History (Botany Series.)
8(3): 227–332.
126. Anonymous (1991). Natural Resources of Sri Lanka:
Conditions and Trends. pp. 280. Natural Resources,
Energy and Science Authority of Sri Lanka, 47/5,
Maitland Place, Colombo 07.
127. Miththapala S., Seidensticke J., Coomaraswamy
U., & Kumarasingham S., O’Brien S. J. (1996).
Phylogeographic subspecies recognition in leopards
(Panthera pardus): molecular genetic variation.
Conservation Biology 10 (4):1115-1132
128. Uphyrkina O., Johnson W E., Quigley H., Miquelle
D. M. L., Bush M. & O’ Brien S. J. (2001). Phylogenetics,
genome diversity and origin of modern leopard, Panthera
pardus. Molecular Ecology 10: 2617-2633.
129. Fernando P., Pfrender M. E., Encalada S. E & Lande,
R. (2000). Mitochondrial DNA variation, phylogeography
and population structure of the Asian elephant. Heredity
84(3): 362-372.
Nimal Gunatilleke et. al
60
Journal of the National Science Foundation of Sri Lanka 36 Special Issue
November 2008
130 Gower D.J., Bahir M.M., Mapatuna Y., Pethiyagoda
R., Raheem D. & Wilkinson M. (2004). Molecular
phylogeny of Sri Lankan ichthyophis (Amphibia:
Gymnophiona:Ichthyophiidae) with discovery of a cryptic
species. Raffles Bulletin of Zoology Supplement No.
12: 153-161.
131 Murawski D.A., Dayanandan B. & Bawa K.S. (1994a).
Outcrossing rates of two endemic Shorea species from
Sri Lankan tropical rain forests. Biotropica 26 : 23-29.
132 Murawski D.A., Gunatilleke I.A.U.N. & Bawa K.S.
(1994c). The effects of selective logging on inbreeding
in Shorea megistophylla (Dipterocarpaceae) from
Sri Lanka. Conservation Biology 8: 997-1002.
133 Murawski D.A. & K.S. Bawa (1994b). Genetic structure
and mating system of Stemonoporus oblongifolius
(Dipterocarpaceae) in Sri Lanka. American Journal of
Botany 81:155-160.
134 Gamage D.T., Silva M.P., Yoshida A., Szmidt A.E.
& Yamazaki T. (2003). Molecular phylogeny of
Sri Lankan Dipterocarpaceae in relation to other Asian
Dipterocarpaceae based on chloroplast DNA sequences.
Tropics 13: 79-87.
135 Stacy E.A. (2001). Cross-fertility in two tropical tree
species: evidence of inbreeding depression within
populations and genetic divergence among populations.
American Journal of Botany 88: 1041-1051.
136 Ratnayake R.M.C.S., Su Y.C.F., Gunatilleke I.A.U.N.,
Wijesundera D.S.A. & Saunders R.M.K. (2006).
Reproductive biology of two sympatric species of
Polyalthia (Annonaceae) in Sri Lanka II. Breeding
systems and population genetic structure. International
Journal of Plant Science 167: 495-502.
137 Anonymous (2007). FAO/Government Co-operative
Programme: Final Report on the establishment of the
National Information Sharing Mechanism (NISM) on
the Implementation of the Global Plan of Action for the
Conservation and Utilization of Plant genetic Resources
for Food and Agriculture in Sri Lanka (Eds. D.H.
Muthukuda Arachchi, C.S. Weeraratna & L. Amarasinghe)
AG: GCP/RAS/186/JPN Field Document No. 2007/03.
pp. 189.
138 Wijesinghe L.C.A. de S., Gunatilleke I.A.U.N.,
Jayawardana S.D.G., Kotagama S. W., Gunatilleke,
C.V.S. (1993). Biological Conservation in Sri Lanka: A
National Status Report. The World Conservation Union,
53 Horton Place, Colombo 07.
139 Anonymous (1996). An investment proposal for the
conservation and sustainable use of medicinal plants in
Sri Lanka IUCN Sri Lanka, Colombo.
140 Myres N., Mittermeier R. A., Mittermeier C., Da Fonseca
G. A. R. & Kent J. (2000). Biodiversity hotspots for
conservation priorities. Nature 403: 853-858.
141 Olson S., Sadacharan D., Samarakoon J.I., White A.T.,
Wickramaratne H.J.M. & Wijeratne M.S. (1992). Coastal
2000: Recommendations for a Resource Management
Strategy for Sri Lanka’s Coastal Region. Coast
Conservation Department & Coastal Resources Center,
The University of Rhode Island, USA.
142 Rajasuriya A. (2002). Status Report on the condition of
reef habitats in Sri Lanka. In: Coral Reef Degradation
in the Indian Ocean Status Report.(Eds. O. Lindon,
D. Souter, D. Wilhelmsson & D. Obura) pp. 139-148
CORDIO, University of Kalmar, Sweden.
143 Lindon O., Souter D., Wilhelmsson D. & Obura
D. (2002). Coral Reef Degradation in the Indian Ocean
Status Report. CORDIO, University of Kalmar, Sweden.
144 Fernando P., Wikramanayake E.D. & Pastorini J. (2006).
Impact of tsunami on terrestrial ecosystems of Yala
National Park, Sri Lanka. Current Science 90: 1531-
1534.
145 Perera N. P. (1984). Natural resources, settlements and
land use. In: Ecology and Biogeography of Sri Lanka.
(Ed. C.H. Fernando) pp. 453 - 492.
146 Pethiyagoda R. (2007). Pearls, Spices and Green Gold:
An Illustrated History of Biodiversity Exploration in Sri
Lanka. pp. 241. Willife Heritage Trust, Publications Pvt.
Ltd., Colombo.
147 Karunaratne N. (1987). Forest Conservation in
Sri Lanka from British Colonial Times 1818 - 1982. pp.
101. Trumpet Publishers Pvt Ltd., Colombo.
148 Gunatilleke C.V.S. Gunatilleke I.A.U.N. (1983). A forestry
case study of the Sinharaja rain forest in Sri Lanka. In:
Forest and Watershed Development and Conservation
in Asia and the Pacific. (Ed. L.Hamilton) pp. 289-358.
Westview Press, Colorado, USA
149 Perera G.A.D. (2001). The secondary forest situation in
Sri Lanka: a review. Journal of Tropical Forest Science
13(4): 768-785.
150 Jayaweera D.M.A. (1981-1982). Medicinal Plants Used
in Ceylon. Parts 1-5. National Science Council, 47/5,
Maitland Place, Colombo 07.
151 McDermott M., Gunatilleke C.V.S. & Gunatilleke
I.A.U.N. (1990a). The Sinharaja rain forest: conserving
both biological diversity and a way of life. The Sri Lanka
Forester 19 (3&4): 3-22.
152 Mc Dermott, M., Gunatilleke C.V.S. & Gunatilleke
I.A.U.N. (1990b). The Sinharaja rain forest: the
compatibility of biological and cultural conservation. The
Sri Lanka Forester 22: 3-28.
153 Gunatilleke I.A.U.N. & Gunatilleke C.V.S. (1993).
Underutilized food plant resources of Sinharaja rain
forest in Sri Lanka. In: Food and Nutrition in the Tropical
Rain Forest: Bicultural Interactions, (C.M. Hladik, A.
Hladik, H. Pagazy. O.F. Linaries & M. Hadley). Man and
Biosphere Series, Vol.15, UNESCO, Paris and Parthenon
Publishing, Carnforth, UK. pp. 183-198
154 de Zoysa N. & Vivekanandan K. (1994). Rattans of
Sri Lanka: an illustrated !eld guide pp. 82. Forest
Department, Sri Lanka.
155 Ashton, Mark S., Gunatilleke Savitri., De Zoysa Neela.,
Gunatilleke Nimal., Dassanayake M.D. & Wijesundera
Siril (1997). A Field Guide to the Common Trees and
Shrubs of Sri Lanka. pp. 431. The Wildlife Heritage Trust
of Sri Lanka.
156 Rajapakse L.K., Padmalal U.K.G.A. & Kotagama
S.W. (2002). Impact of sambar deer (Cervus unicolor
Biodiversity of Sri Lanka 61
Journal of the National Science Foundation of Sri Lanka 36 Special Issue November 2008
unicolor) on the vegetation at the Horton Plains
National Park. Proceedings of the eighth annual
forestry and environment symposium 2002, Dept. of
Forestry and Environmental Science, University of Sri
Jayawardenepura, Gangodawila, Nugegoda.
157 Padmalal U.K.G.A., Takatsuki S. & Jayasekara P. (year)
Food habits of sambar Cervus unicolor at the Horton
Plains National Park, Sri Lanka. Ecological Research
18(6): 775 - 782.
158 Gunawardana E.R.N., Rajapakse U., Nanadasena
K.A., Rosier P.T.W Chandrasiri N. (1998) Water quality
use in the uplands of Sri Lanka, (Ed. H.P.M. Gunasena)
In: Proceedings of the Final Workshop University of
Peradeniya-Oxford Forestry Institute Link Project.
pp. 37-44.
159 Anonymous (1999). Biodiversity Conservation in Sri
Lanka: A Framework for Action. pp.126. Ministry of
Forestry and Environment, Battaramulla.
160 Ekaratne K., Fernando R.H.S.S., de Silva S.,
Bambaradeniya C. M. B. & de Silva D. (2003). A
Comparison of the Conservation and Legal Status of the
Fauna and Flora of Sri Lanka. pp. 163. IUCN Sri Lanka,
Colombo.
161 Anonymous (2003). Caring for the Environment 2003 -
2007: Path to Sustainable Development pp. 152. Ministry
of Environment and Natural Resources, Battaramulla .
162 Anonymous (2007). The UN Decade of Education for
Sustainable Development (DESD 2005 - 2014) the First
Two Years. Pp. 64. UNESCO, France.
163. UNESCO (2007). Biodiversity in UNESCO, pp. 32.
UNESCO, Paris.