Editor

Craig Hassapakis

Berkeley, California, USA

Associate Editors

Raul E. Diaz Howard O. Clark, Jr. Erik R. Wild

University of Kansas, USA Garcia and Associates, USA University of Wisconsin-Stevens Point, USA

Assistant Editors

Alison R. Davis

University of California, Berkeley, USA

Daniel D. Fogell

Southeastern Community College, USA

Editorial Review Board

David C. Blackburn

California Academy of Sciences, USA

Bill Branch

Port Elizabeth Museum, SOUTH AFRICA

Jelka Crnobrnja-Isailovc

IBISS University of Belgrade, SERBIA

C. Kenneth Dodd, Jr.

University of Florida, USA

Lee a. Fitzgerald

Texas A&M University, USA

Adel A. Ibrahim

Ha’il University, SAUDIA ARABIA

Harvey B. Lillywhite

University of Florida, USA

Julian C. Lee

Taos, New Mexico, USA

Rafaqat Masroor

Pakistan Museum of Natural History, PAKISTAN

Peter V. Lindeman

Edinboro University of Pennsylvania, USA

Jaime E. Pefaur

Universidad de Los Andes, VENEZUELA

Jodi J. L. Rowley

Australian Museum, AUSTRALIA

Henry R. Mushinsky

University of South Florida, USA

Rohan Pethiyagoda

Australian Museum, AUSTRALIA

Peter Uetz

Virginia Commonwealth University, USA

Elnaz Najafimajd

Ege University, TURKEY

Nasrullah Rastegar-Pouyani

Razi University, IRAN

Larry David Wilson

Instituto Regional de Biodiversidad, USA

Allison C. Alberts

Zoological Society of San Diego, USA

Michael B. Eisen

Public Library of Science, USA

Russell A. Mittermeier

Conservation International, USA

Advisory Board

Aaron M. Bauer

Villanova University, USA

James Hanken

Harvard University, USA

Robert W. Murphy

Royal Ontario Museum, CANADA

Walter R. Erdelen

UNESCO, FRANCE

Roy W. McDiarmid

USGS Patuxent Wildlife Research Center, USA

Eric R. Pianka

University of Texas, Austin, USA

Antonio W. Salas

Environment and Sustainable Development, PERU

Dawn S. Wilson

AMNH Southwestern Research Station, USA

Honorary Members

Carl C. Gans Joseph T. Collins

(1923-2009) (1939-2012)

Cover :

Green Pit-viper Trimeresurus trigonocephalus eaptured in Lakegala, Dumbara Hills, Knuekles World Heritage site, Sri Lanka. The sole rep-

resentative of the genus Trimeresurus on the island of Sri Lanka; an endemie speeies. Noeturnal, sluggish and arboreal this snake is found in

forested areas and oeeasionally in well-wooded home gardens and plantations sueh as tea, eoffee, eardamom, eoeoa, and elove nutmeg. More

eommonly distributed in the wet zone of the eountry but also found in the dry zone as well. Commonly found on low bushes and deseending to

the ground to seareh for prey at night. Generally found elose to streams. Photo Imesh Numan Bandura.

Amphibian & Reptile Conservation — Worldwide Community- Supported Herpetological Conservation (ISSN: 1083-446X; elSSN: 1525-9153) is

published by Craig Hassapakis/Amp/z/Z^/tm & Reptile Conservation as full issues at least twiee yearly (semi-annually or more often depending on

needs) and papers are immediately released as they are finished on our website; http://amphibian-reptile-conservation.org; email:

are.publisher@gmail.eom

Amphibian & Reptile Conservation is published as an open access journal. Please visit the official journal website at:

http://amphibian-reptile-eonservation.org

Instruetions to Authors : Amphibian & Reptile Conservation aeeepts manuseripts on the biology of amphibians and reptiles, with emphasis on

eonservation, sustainable management, and biodiversity. Topies in these areas ean inelude: taxonomy and phylogeny, speeies inventories, distri-

bution, conservation, species profiles, ecology, natural history, sustainable management, conservation breeding, citizen science, social network-

ing, and any other topie that lends to the eonservation of amphibians and reptiles worldwide. Prior eonsultation with editors is suggested and

important if you have any questions and/or eoneerns about submissions. Further details on the submission of a manuseript ean best be obtained

by eonsulting a eurrent published paper from the journal and/or by aeeessing Instruetions for Authors at the Amphibian and Reptile Conservation

website: http://amphibian-reptile-eonservation.org/submissions.html

Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium,

provided the original author and source are credited.

Amphibian & Reptiie Conservation 5(2):1-13.

The herpetofauna of a small and unprotected patch of

tropical rainforest in Morningside, Sri Lanka

' 3RETER JANZEN AND ^MALAKA BOPAGE

^Rheinallee 13, 47119 Duisburg, GERMANY ^Biodiversity Education & Exploration Society (BEES) 63/c Wackvella road Galle 80000, SRI LANKA

Abstract . — Morningside is an exceptional area in Sri Lanka with highly endemic herpetofauna. How-

ever, this relictual forest area lies inside a tea plantation and is mostly lacking conservation protec-

tion. Species inventories of remaining rainforest patches are currently incomplete, and information

about the behavior and ecology of the herpetofauna of Morningside is poorly known. In our survey,

we identified 13 amphibian species and recorded an additional two species that could not be identi-

fied with existing keys. We determined 11 reptile species from this patch of forest, and another un-

identified Cnemasp/s gecko was recorded. We did not assess the herpetofauna outside of this forest

patch. Some species are described for the first time in Morningside, suggesting a wider distribution

in Sri Lanka. We also document a call from a male Pseudophiiautus cavirostris for the first time.

Perspectives for future surveys are given.

Key words. Survey, Morningside, Sri Lanka, herpetofauna, conservation, Pseudophiiautus cavirostris

Citation: Jansen, P. and Bopage, M. 2011 . The herpetofauna of a small and unprotected patch of tropical rainforest in Morningside, Sri Lanka. Amphib-

ian & Reptile Conservation 5(2) :1 -1 3(e26).

Introduction

Sri Lanka is a small (65,610 km^) island south of India.

The island lies between latitudes 5°55’ and 9°51’ N and

longitudes 79°41’ and 81°54’ E. Sri Lanka is divided into

four different climatic zones (Domroes and Roth 1998):

dry, wet, transitional, and semiarid. The dry zone is situ-

ated in the eastern and northern parts of the island, cover-

ing 60% of the total land area. Annual rainfall is between

1250 and 1900 mm, and the mean annual temperature

ranges from 27° to 30° C. Floristically, the dry zone is

characterized by monsoon forests and thorn scrublands.

The wet zone encompasses southwestern Sri Lanka, cov-

ering 23% of the total land area and receiving an annual

rainfall of 2500-5000 mm. The natural vegetation con-

sists of evergreen, semi-evergreen, and rain forest. Be-

tween these two zones lies an intermediate transitional

zone, with annual rainfall between 1900 and 2500 mm.

The two semiarid zones (in the southeast and northwest)

receive less than 1250 mm of rainfall annually. Within

these zones, climate can also vary along elevational gra-

dients. In mountainous regions, the temperature is lower

and can approach freezing at times. This high elevation

climate has been recognized previously from both the

Central Mountains and the Knuckles Mountains, and

more recently from the Rakwana Hills. All three of these

mountainous regions have a different climate from the

surrounding area, as expected (Werner 2001). The Morn-

ingside area lies in the Rakwana Hills.

Correspondence. ^ Email: pjanzen@ gmx.de

In our attempt to understand the biodiversity of Sri

Lanka, scientists from the Wildlife Heritage Trust (WHT)

have made great progress in naming many new species

and significantly expanding our knowledge of the region.

However, there are likely still undescribed amphibians

and reptiles in Sri Lanka (Anslem de Silva, pers. comm.,

Krvavac, pers. comm). Due to the high levels of ende-

mism found in Morningside, scientists and conservation

organizations like Conservation International have iden-

tified it as a region of high conservation priority. Located

in the eastern part of the Sinharaja forest, Morningside

has also been declared a Man and Biosphere Reserve

(MAB Reserve) under the UNESCO World Heritage

Convention. Sinharaja is the largest remaining tropical

rainforest in Sri Eanka, but most unprotected parts of the

forest in Morningside are logged. Today, only a few for-

est fragments remain.

Methodology

To survey Morningside for reptiles and amphibians, field-

work was conducted for three days and nights in a small

patch of remaining forest near the town of Suriyakanda in

July 2010. This patch of forest lies inside a tea plantation

and lacks any conservation protection, and it is possible

that it will be cleared for tea plants in the near future. The

coordinates of our survey starting point were identified

amphibian-reptile-conservation.org

001

October 2011 I Volume 5 I Number 2 I e26

Janzen and Bopage

with a handheld GPS (Garmin eTrex) as 6° 27’ 17” N

and 80° 37’ 9” E at an elevation of 975 m asl (above sea

level). We could not ascertain the size of the forest patch

using the available resources. The forest lacks large trees

(above 10 m) and the canopy is not completely closed. In

this open canopy, sufficient light reached the ground and

bushes were able to grow; it was often possible to see the

sky through holes in the canopy. No attempts were made

to identify vegetation. No rain was recorded during the

study period, but strong winds prevailed during most of

the sampling time. The surveys were conducted by walk-

ing along trails and a stream that fiows through the forest,

as well as by searching in and around ponds. The ponds

had a depth of less than 60 cm and were considered to

be temporary. Dead logs and rocks were overturned and

leaf litter was checked for reptiles and amphibians. These

surveys were done during daytime and at night between

8 p.m. and midnight.

Results

During the field trips, we found 15 species of amphib-

ians, although two of these were unidentifiable using

current taxonomy keys (not listed below). A total of 11

species of reptiles were identified, plus one unidentified

gecko. All identified species are listed in Table 1.

Reptiles

Gekkonidae

Cnemaspis sp.

The genus Cnemaspis consists of day-active geckos. The

species are more or less brownish to grayish in color-

ation. We found all specimens inside or around a small

house nearby the forest. The geckos are common around

the house, and they lay eggs in small holes in the door-

frame. We could not find evidence for communal egg lay-

ing. This behavior is described for another member of the

Cnemaspis sp.

genus Cnemaspis, and we found a communal laying site

of Cnemaspis at Morningside Estate, only a few kilome-

ters away from this forest patch. Species identification of

these specimens was not possible, as this genus must be

reviewed for the whole of Sri Eanka, and in particular for

Morningside. Several new species have been discovered,

but remain undescribed (Anslem de Silva, pers. comm.).

Cyrtodactylus subsolanus

This gecko formerly belonged to the species C. fraenatus

and was identified as a distinct species in by Batuwita

and Bahir (2005). We found an adult specimen with to-

tal length 20 cm inside the house foraging for insects at

night and a single young specimen in a bush during a

trip in the late evening. The day gecko C. subsolanus is

restricted to Morningside.

Cyrtodactylus subsolanus.

amphibian-reptile-conservation.org

002

October 2011 I Volume 5 I Number 2 I e26

Herpetofauna of Morningside, Sri Lanka

Tropical rainforest survey area in Morningside, Sri Lanka.

Table 1 . Checklist of amphibians and reptiles found during the survey

Amphibians

Reptiles

Bufonidae

Agamidae

Adenomus kelaartii {GunVner, 1858) endangered*

Calotes calotes (Linnaeus, 1758) near threatened

Calotes liolepis Boulenger, 1 885 vulnerable*

Dicroglossidae

Lyriocephalus scutatus (Linnaeus, 1758) near threatened*

Fejervarya kirtisinghei (Manamendra-Arachchi and Gabadage,

Otocryptis wiegmanni Wag\er, 1830 near threatened*

1996) least concern*

Gekkonidae

Microhylidae

Cnemaspis spec.

Ramanella obscura (Gunther, 1864) near threatened*

Cyrtodactylus subsolanus Batuwita and Bahir, 2005 not

evaluated*

Ranidae

Geckoella triedrus (Gunther, 1864) near threatened*

Hylarana temporalis (Gunther, 1864) near threatened

Scincidae

Rhacophoridae

Lankascincus taprobanensis (Kelaart, 1854) near threatened*

Pseudophilautus cavirostris (Gunther, 1869 ) endangered*

Pseudophilautus fergusonianus (Ahl, 1927) least concern*

Pseudophilautus folicola (Manamendra-Arachchi and Pethiya

goda 2005) endangered*

Pseudophilautus procax (Manamendra-Arachchi and Pethiya

Colubridae

Ahaetulla nasuta (Bonnaterre, 1790)

Dendrelaphis pictus (Gmelin, 1 789)

Viperidae

goda 2005) critically endangered*

Pseudophilautus reticulatus (Gunther, 1869) endangered*

Hypnale hypnale (Laurenti, 1768)*

Pseudophilautus singu (Meegaskumbura, Manamendra-Arach

chi and Pethiyagoda 2009) not evaluated*

Pseudophilautus stictomerus (Gunther, 1876) near threatened*

Polypedates cruciger Blyth, 1 852 least concern*

Polypedates fastigo Manamendra-Arachchi and Pethiyagoda

Trimeresurus trigonocephalus (Latreille, 1801) vulnerable*

2001 critcally endangered*

*Asterisk stands for endemic to Sri Lanka

amphibian-reptile-conservation.org

003

October 2011 | Volume 5 | Number 2 | e26

Janzen and Bopage

Geckoella triedrus

This small gecko is a typical inhabitant of forests in the

wet zone, but it is recorded from some parts of the dry

zone as well. Das and De Silva (2005) restricted the el-

evational distribution to 700 m asl. However, we found

our only specimen active at night at an elevation of 975

m asl. Geckoella triedrus is a small brown to black col-

ored gecko with tiny whitish dots on the dorsum. This

gecko is a member of the leaf litter herpetofauna living

on the ground, and it is difficult to find.

Geckoella triedrus.

Agamidae

Calotes calotes

Calotes calotes is a widespread arboreal agamid found

all over Sri Lanka up to 1500 m asl. The distribution

ranges north into India. This agamid lizard is a typical

anthropophilic species and is often found in gardens. We

found a male C. calotes sleeping in the late evening at

the forest border.

Calotes calotes.

a slightly higher rainfall than the surrounding area. It is

distributed in forests and plantations up to 1000 m asl.

Our detection of C. liolepis in Morningside represents

the highest regions in the distribution. Calotes liolepis is

endemic to the region. This agamid species is difficult to

find because it climbs the stems of trees and then curls

around the stem, avoiding detection. All three specimens

(one female and two males) that we found sat on a stem

at heights between 4 and 6 m. One of the males had two

bluish stripes laterally and an orange throat. The female

was grayish colored. Somaweera found a specimen with

red stripes (Manthey 2008). One of the authors (M.B.)

found C. desilvai on an earlier trip in this forest patch.

Calotes desilvai looks quite similar to C. liolepis and is

restricted to a small part of the Morningside area (Ba-

hir and Maduwage 2005). This is one of the few places

where both species live in sympatry. However, we did

not detect any C. desilvai on this trip.

Calotes liolepis.

Otocryptis wiegmanni

Calotes liolepis

This agamid lizard is generally restricted to the wet

zone, with a few exceptions in the intermediate and dry

zone. In these drier areas, it is found on small hills with

The kangaroo lizard is very common in the forests of

Morningside. We found adults and young specimens fre-

quently. This agamid is distributed throughout the wet

zone and some parts of the intermediate zone as well.

Only one species of the genus was described for Sri Lan-

ka until Bahir and Silva (2005) described a new species

amphibian-reptile-conservation.org

004

October 2011 I Volume 5 I Number 2 I e26

Herpetofauna of Morningside, Sri Lanka

(O. nigristigma). Otocryptis nigristigma is restricted to

the dry and intermediate zones. Male O. wiegmanni have

a black patch on the dewlap, and by this they can be dis-

tinguished from O. nigristigma. Otocryptis wiegmanni is

able to run bipedally when fleeing. Otocryptis wiegman-

ni can be found active during daytime or sleeping in the

darkness on branches of trees and bushes.

Otocryptis wiegmanni male specimen.

Otocryptis wiegmanni s\eep\ng.

Lyriocephalus scutatus

Lyriocephalus scutatus is restricted to the wet zone and

few places of the intermediate zone below 1600 m asl,

Lyriocephalus scutatus young specimen.

where it inhabits forests and home gardens. It is a slow-

moving species and is mostly arboreal. Most specimens

are light green or yellowish in coloration, although fe-

males are sometimes grayish or brownish. Young spec-

imens are brownish and live on or near the ground in

bushes or small trees. A unique defensive posture of this

species is the display of the deep red color of the mouth.

Lyriocephalus scutatus can easily be found in the dark-

ness when they sleep and hang on tree stems. In the light

of a torch, one can see them easily by the light color-

ation of the body. We found L. scutatus often, from very

young to adult male specimens during both daytime and

at night.

Scincidae

Lankascincus taprobanensis

Lankascincus are ground living species found in leaf lit-

ter. It is difficult to photograph these skinks because they

quickly hide under leaf litter upon detection. Lankascin-

cus taprobanensis is a mountainous species, distributed

from 1000 m to 2300 m asl. We found this skink at their

lowest distribution level in Morningside. The skinks are

active during daytime and can be easily photographed at

night.

amphibian-reptile-conservation.org

005

October 2011 | Volume 5 | Number 2 | e26

Janzen and Bopage

Lankascincus taprobanensis.

Hypnale zara.

Colubridae

Ahaetulla nasuta

Only one specimen was found in tree branches at the bor-

der of the forest at night. Ahaetulla nasuta is widely dis-

tributed across Sri Lanka and mainland Asia. This snake

is often found in gardens in every climatic zone. There

are no color varieties of A. nasuta in Sri Lanka. This

opistoglyph snake is green-colored and becomes mottled

when disturbed.

Dendrelaphis tristis

This slender and long snake has nearly the same distribu-

tion as A. nasuta, and we found one specimen nearly at

the same place as the A. nasuta specimen. Dendrelaphis

tristis is a common snake, more typically found in the

lower parts of Sri Lanka. Das and De Silva (2005) gave a

distribution range up to 750 m asl. We found this species

200 m higher in Morningside. The snake was hiding in

bushes at night.

Viperidae

Hypnale zara

This venomous snake is endemic to Sri Lanka. It is a

small brownish snake found in mountain and submon-

tane forests living in leaf litter, where it can easily be

overlooked. We found a specimen hiding around a pond

at night.

Trimeresurus trigonocephalus

Trimeresurus trigonocephalus is an arboreal snake with

greenish ground color and often variegated black pat-

terns. This species is distributed throughout Sri Lanka

below 1075 m asl. We found one specimen hanging on

branches next to a pond in the dark. It is a very docile

species; the snake did not try to bite, but it did try to

escape.

Trimeresurus trigonocephalus.

amphibian-reptile-conservation.org

006

October 2011 I Volume 5 I Number 2 I e26

Herpetofauna of Morningside, Sri Lanka

Ramanella obscura.

Adenomus kelaartii.

Dicroglossidae

Fejervarya kirtisinghei

Trimeresurus trigonocephalus.

Amphibians

Bufonidae

Adenomus kelaartii

Adenomus kelaartii is a small slender toad found near

streams, which is where we found our only specimen

during the survey. It is a ground-dwelling species, but

it can sometimes be found climbing on trees. Adenomus

kelaartii is restricted to the wet zone and mountainous

areas of Sri Lanka. There are no descriptions of eggs or

tadpoles in nature, but there is a description of tadpoles

from captive bred specimens (Haas et al. 1997; Haas

1999). We found one specimen together with Hylarana

temporalis.

Fejervarya kirtisinghei.

Microhylidae

Ramanella obscura

Ramanella obscura is a small species (32 mm) living on

the ground in leaf litter in shaded forests, but it some-

times climbs on trees and can be found in tree holes up to

two meters high. It is distributed throughout the wet zone

up to 1200 m asl. We found several specimens near or in-

side ponds. Egg clutches rest in a single layer on the wa-

ter surface. We found R. obscura tadpoles together with

tadpoles of Fejervarya kirtisinghei in the pond. Breeding

of R. obscura in phytotelmata is described, but we only

found egg clutches in ponds.

This ranid like species is widely distributed in the low-

land areas of Sri Lanka in the wet and the dry zone. In the

past, F. kirtisinghei has been confused with F. greeni. The

latter is restricted to the higher elevations of Sri Lanka.

We found F. kirtisinghei near ponds together with Hyla-

rana temporalis and Ramanella obscura. We observed

tadpoles with the typical black tag in the pond.

Ramanella obscura egg masses.

amphibian-reptile-conservation.org

007

October 2011 I Volume 5 I Number 2 I e26

Janzen and Bopage

Ramanella obscura tadpoles in pond.

Ranidae

Hylarana temporalis

This is a typical species of the forest patch in Morning-

side. It is widely distributed in Sri Lanka’s wet zone from

the lowlands up to 1800 m asl. The frogs are mostly

brownish-colored, with cross bars on the arms and legs.

We found H. temporalis near the stream and near ponds,

where the ground is wet or muddy. One frog had only

one hind foot.

Hylarana temporalis.

Hylarana temporalis with missing foot.

Rhacophoridae

Pseudophilautus cavirostris

An arboreal species, P. cavirostris is perhaps found most

often in canopies (Dutta and Manamendra-Arachchi

1996). This frog reaches 50 mm in length and has a tu-

berculated dorsum and fringes along the lower arms and

tarsus. The coloration can be greenish or mottled with

grey and brown. The frog is well camouflaged to look

like lichens on a stem and is difficult and rare to And.

Descriptions of eggs and mating behavior are not giv-

en elsewhere. We found a male specimen calling from

leaves 1.5 m above ground around 11 p.m. Manamendra-

Arachchi and Pethiyagoda (2005) suggested that males

do not come down from the canopy because they could

not And male specimens.

Pseudophilautus cavirostris calling.

Pseudophilautus cavirostris.

amphibian-reptile-conservation.org

008

October 2011 I Volume 5 I Number 2 I e26

Herpetofauna of Morningside, Sri Lanka

Pseudophilautus fergusonianus

Pseudophilautus procax

This frog is found on trees and rocks in rainforests and

rubber plantations in the hills of the wet zone between

300 and 700 m asl (Manamendra-Arachchi and Pethiya-

goda 2005). We found several specimens, but only in-

side or at the house where we also found Cnemaspis. No

specimens were observed in the forest. The coloration of

P. fergusonianus gave an ideal camouflage on the house

walls. This frog reaches 45 mm (females).

Pseudophilautus fergusonianus.

Pseudophilautus folicola

Pseudophilautus folicola was described as a lowland

species from the wet zone (Manamendra-Arachchi and

Pethiyagoda 2009). Our survey expands the distribution

up to 975 m asl. It seems to be a common species, even

found hiding in the daytime on garden plants.

Pseudophilautus folicola.

Pseudophilautus procax is a tiny species (27 mm) found

at night on leaves one to two meters above the ground.

The coloration is light brown, sometimes a bit yellowish,

with a yellowish to white infraorbital patch and red fin-

gertips. This species is endemic to Morningside.

Pseudophilautus procax.

Pseudophilautus reticulatus

Pseudophilautus reticulatus is a larger species of the ge-

nus, with females reaching 61 mm. The scientific name

for this species is derived from the markings down the

lateral sides of the body and on the inner part of the

femora. It is an arboreal species that comes down from

canopies at night. In our estimation, this frog should be

distributed in forests of the wet zone up to an elevation of

975 m asl. The true distribution of this species is unclear.

Pseudophilautus reticulatus: note markings down the lat-

eral sides of the body and on the inner part of the femora.

amphibian-reptile-conservation.org

009

October 2011 | Volume 5 | Number 2 | e26

Janzen and Bopage

Pseudophilautus reticulatus.

Pseudophilautus singu

We found specimens with grayish or light brownish

ground coloration, which is in contrast to the original de-

scription of the species (Meegaskumbura, Manamendra-

Arachchi, and Pethiyagoda 2009). It is a small species

(males less than 20 mm), but females are not described

and their size is unknown and undescribed in scientific

papers. Pseudophilautus singu was found near ponds on

leaves 1-2 m above the ground.

Pseudophilautus singu.

Pseudophilautus stictomerus

Pseudophilautus stictomerus is a small species (23 to

36 mm) from Sri Lanka’s wet zone. Although it was as-

sumed that this species is distributed to 700 m asl, we

found this species at an elevation of 975 m asl. We found

a small specimen, brownish-colored, with a fine white

line from snout to vent and further along the hind legs

and a yellow throat. The coloration of the throat could be

an indicator for a male specimen.

Pseudophilautus stictomerus.

Polypedates cruciger

Polypedates cruciger is a large rhacophorid frog (male 60

mm and female 90 mm). It is a common species, found

from the wet zone to the dry zone. It is a species that

can be found in gardens and inside houses. Mating and

breeding of this species is well known and documented

(Herrmann 1993). We found two specimens at a pond in-

side the forest, sympatric with Tarugafastigo.

Polypedates cruciger.

amphibian-reptile-conservation.org

010

October 2011 I Volume 5 I Number 2 I e26

Herpetofauna of Morningside, Sri Lanka

Polypedates cruciger.

Taruga fastigo

Taruga fastigo is a beautiful tree frog and very similar

to P. longinasus. Taruga fastigo is restricted to Morning-

side, and P. longinasus is a lowland species in forests of

the wet zone. Unfortunately, there is no genetic verifica-

tion that these are separate species. However, it is pos-

sible that both species live sympatrically in the Sinharaja

forest. Taruga fastigo is a common species in this forest

patch, and we found young and adult frogs at night on

leaves and branches up to 2 m above ground. At the pond,

we found a foam nest of Taruga fastigo containing a few

unfertilized eggs. Further observations of Taruga fastigo

are necessary, especially for breeding information, be-

cause this is a critically endangered species.

Polypedates fastigo.

Taruga fastigo.

Taruga fastigo foam nest.

Discussion

During our brief survey, we found an interesting diver-

sity of reptile and amphibian species, some of which

were previously unknown from Morningside. This sur-

vey shows how much knowledge we are lacking about

the distribution and ecology of reptiles and especially of

the amphibians of Sri Lanka. Further investigations are

necessary to answer these and future questions. The be-

havior and ecology of some of these species is currently

not well known. One example of this lack of knowledge

amphibian-reptile-conservation.org 011 October 2011 | Volume 5 | Number 2 | e26

Janzen and Bopage

is that we provide the first published record of a calling

male P. cavoristris. This small patch of remaining tropi-

cal rainforest is ecologically valuable, an ideal place for

a larger study of the ecology of such small forest patch-

es and also for the ecology of these species of reptiles

and amphibians. Also, little is known about the mating

behavior and breeding of Sri Lankan amphibians (Ka-

runarathna and Amarasinghe 2007). Future research is

necessary and should be done in both nature and in cap-

tivity, as was previously conducted by Wildlife Heritage

Trust at Agrapatana (Bahir et al. 2005).

This survey also highlights the need for more re-

search at Morningside because some expected species

were not detected on our trip. We could not find any

specimens of the genus Ceratophora (C. erdeleni and C.

karu), even though the Morningside Estate where they

are known to occur is not far away from this forest patch.

Both species are restricted to the Morningside region. We

also found a few frog species only at Morningside Estate

{Pseudophilautus poppiae, P sordidus, and P decoris),

but not in the forest patch. It is possible that these frogs

could be present in the forest patch as well, but escaped

detection. One of the authors (M. B.) found Microhyla

karunaratnei on a previous trip, but we did not find any

specimens on the trip described here. We also found two

species of PseudophUautus that we could not accurately

identify to the species level. These uncertainties, as well

as its conservation priority, suggest that Morningside

should be a target for future research on reptiles and am-

phibians.

Acknowledgments. — The authors thank Rohan Pethi-

yagoda for reviewing the article and Craig Hassapakis

for publishing this paper.

References

Bahir, M. M. and Maduwage, K. P. 2005. Calotes desil-

vei, a new species of agamid lizard from Morningside

Forest, Sri Lanka. Raffles Bulletin of Zoology, Supple-

ment 12:381-392.

Bahir, M. M., Meegaskumbura, M., Manamendra-Arach-

chi, K., Schneider, C. J., and Pethiyagoda, R. 2005.

Reproduction and terrestrial direct development in

Sri Lankan Shrub-Frogs (Ranidae: Rhacophorinae:

Philautus). Raffles Bulletin of Zoology, Supplement

12:339-350.

Bahir, M. M. and Silva, A. 2005. Otocryptis nigristigma,

a new species of agamid lizard from Sri Lanka. Raffles

Bulletin of Zoology, Supplement 12:393-406.

Das, I. and De Silva, A. 2005. A Photographic Guide to

Snakes and Other Reptiles of Sri Lanka. New Holland

Publishers, London. 144 p.

Deraniyagala, P. E. P. 1939. The Tetrapod Reptiles of

Pseudophilautus unknown species.

Ceylon, Volume 1: Testudinates and Crocodilians .

Museum of Natural History, Colombo. 412 p.

Deraniyagala, P. E. P. 1953. A Colored Atlas of Some Ver-

tebrates from Ceylon, Volume 2: Tetrapod Reptilia.

Ceylon Government Press, Museum of Natural His-

tory, Colombo. 101 p.

Deraniyagala, P. E. P. 1955. A Colored Atlas of Some Ver-

tebrates from Ceylon, Volume 3: Serpentoid Reptilia.

Ceylon Government Press, Museum of Natural His-

tory, Colombo. 121 p.

Domroes, M. and Roth, H. 1998. Sri Lanka - Past and

Present: archaeology, geography, economics: Select-

ed papers on German research. Margraf Publishers

GmbH, Weikersheim. 1997 p.

Dutta, S. K. and Manamendra-Arachchi, K. 1996. The

Amphibian Fauna of Sri Lanka: a systematic review.

Wildlife Heritage Trust, Colombo, Sri Lanka. 230 p.

Haas, W, Lehr, E., and Kohler, G. 1997. The tadpole of

Bufo kelaartii Gunther 1859 from Sri Lanka. Lyrio-

cephalus 3(2):2-6.

Haas, W. 1999. Zur Biologie von Bufo kelaartii Gunther,

1859. Elaphe 7(2): 16-19.

Herrmann, H.-J. 1993. Haltung und Zucht von Polyped-

ates cruciger cruciger Blyth, 1852. Herpetofauna

15(85):31-34.

Karunarathna, D. M. S. S. and Amarasinghe, A. A. T.

amphibian-reptile-conservation.org

012

October 2011 I Volume 5 I Number 2 I e26

Herpetofauna of Morningside, Sri Lanka

2007. Observations on the breeding behavior of

Philautus regius Manamendra-Arachchi and Pethi-

yagoda 2005 (Amphibia: Ranidae: Rhacophoridae)

in Nilgala, Monaragala District in Sri Lanka. Russian

Journal of Herpetology 14(2): 133-136.

Kelaart, E. E, Wijesinghe, P, Pethiyagoda, R., and Man-

amendra-Arachchi, K. 1998. Prodromus Faunae Zey-

lanicae: A Facsimile Reprint of the 1852 and 1854

texts. Wildlife Heritage Trust, Colombo. 342 p.

Maduwage, K., Silva, A., Manamendra-Arachchi, K.,

and Pethiyagoda, R. 2009. A taxonomic revision of

the South Asian hump-nosed pit vipers (Squamata:

Viperidae; Hypnale). Zootaxa 2232:1-28.

Manamendra-Arachchi, K., Batuwita, S., and Pethiya-

goda, R. 2007. A taxonomic revision of the Sri Lank-

an day-geckos (Reptilia: Gekkonidae: Cnemaspis),

with description of a new species from Sri Lanka and

southern India. Zeylanica 7(1):9-122.

Manamendra-Arachchi, K. and Pethiyagoda, R. 2001.

Polypedates fastigo, a new tree frog (Ranidae: Rha-

cophoridae) from Sri Lanka. Journal of South Asian

Natural History 5(2): 191-199.

Manamendra-Arachchi, K. and Pethiyagoda, R. 2005.

The Sri Lankan shrub-frogs of the genus Philautus

Gistel, 1848 (Ranidae: Rhacophorinae), with descrip-

tion of 27 new species. Raffles Bulletin of Zoology,

Supplement 12:163-303.

Manthey, U. 2008. Terralog. Agamen des siidlichen

Asien 1 / TERRALOG: Agamid Lizards of Southern

Asia, Draconinae 1, (TERRALOG 7a). Aqualog Ver-

lag Gmbh, Lrankfurt, Germany. 160 p.

Meegaskumbura, M., Manamendra-Arachchi, K., and

Pethiyagoda, R. 2009. Two new species of shrub frogs

(Rhacophoridae: Philautus) from the lowlands of Sri

Lanka. Zootaxa 2122:51-68.

Somaweera, R. and Somaweera, N. 2009. Lizards of Sri

Lanka, A Colour Guide with Lield Keys. Edition Chi-

maira / Serpent’s Tale NHBD, Lrankfurt, Germany.

304 p.

Werner, W. 2001. Sri Lanka ’s Magnificent Cloud Forests.

Wht Publications, Colombo. 96 p.

Wrickramasinghe L. J. M. and Munindradasa, D. A. I.

2007. Review of the genus Cnemaspis Strauch, 1887

(Sauria: Gekkonidae) in Sri Lanka with the descrip-

tion of five new species. Zootaxa 1490:1-63.

Manuscript received: 29 December 2010

Accepted: 26 April 2011

Published: 29 October 2011

Peter Janzen gained his Diploma at the Heinrich-Heine-

Universitat in Dusseldorf, Germany in 1990. In 1993 he

finished his Ph.D. studying the activities of mitochondrial

enzymes in human diseases at the Institut of Biochemis-

try at Herinrich-Heine-Universitat. Peter has been inter-

ested in herpetology since childhood and is now active

in coordinating amphibian breeding programmes among

zoos and private persons for the DGHT (Deutsch Ge-

sellschaft fur Herpetologie und Terrarienkunde) and VDZ

(Verband Deutscher Zoodirektoren).

Malaka Bopage was a student of Richmond College

Galle Sri Lanka. He left school after passing the G.C.E.

(A/L) examination in 1998. Malaka has been interested

in herpetology since 1993 and has participated in many

conservation and biodiversity research programs in Sri

Lanka. His research interests include reproductive biol-

ogy and ecology of amphibians from Sri Lanka.

amphibian-reptile-conservation.org

013

October 2011 | Volume 5 | Number 2 | e26

mons Attribution License, which pemiits unrestricted use, distribution, and reproduction in any medium, provided

the original author and source are credited.

Amphibian & Reptiie Conservation 5(2): 14-21.

Predator-induced plasticity in tadpoles of

Polypedates cruciger {Anura: Rhacophoridae)

^KRISHAN ARIYASIRI, ^GAYAN BOWATTE, ^UDENI MENIKE, ^SUYAMA MEEGASKUMBURA, AND

1 ^MADHAVA MEEGASKUMBURA

^Department of Zoology, Faculty of Science, University of Peradeniya, SRI LANKA

Abstract . — ^Aquatic tadpoles morphologically respond to presence of predators in various ways.

Depending on the type of predator, tadpoles develop enhanced escape response abilities in accel-

eration, maneuverability, and speed, and these are correlated to suites of morphological characters,

such as wider, longer, and robust tail related dimensions. Laying eggs away from water, such as in

an arboreal foam nest from which partially developed tadpoles fall into water, could be an adapta-

tion for predator avoidance of eggs and early tadpole stages. Since predation is of concern, even

for these partially developed larvae, we sought to detect predator-induced morphological response

(if any) of these forms compared to fully aquatic tadpoles. We exposed the tadpoles of foam-nesting

Poiypedates cruciger to a natural fish predator, Beiontia signata. We show that at an early (Gosner

stage 29-32) stage, tadpoles exposed to this predator develop a larger body size and increased tail-

length related dimensions.

Key words. Tadpole morphology, plasticity, foam nesting, Polypedates cruciger, predator-induced, morphological

response, amphibian declines

Citation: Ariyasiri K, Bowatte G, Menike U, Meegaskumbura S, Meegaskumbura M. 2011 . Predator-induced plasticity in tadpoles of

Polypedates cruciger {Anura: Rhacophoridae). Amphibian & Reptile Conservation 5(2):14-21(e29).

Introduction

It is well known that aquatic tadpole predators, such as

some dragonfly larvae and fish, induce morphological

changes in aquatic tadpoles (Anderson and Brown 2009;

Buskirk 2002; Teplitsky et al. 2003). Morphological fea-

tures of fully aquatic tadpoles, especially the ones that

are important in swimming, such as tail dimensions, are

known to change in response to predator-type, such as

ambush predators and run-down predators. In the pres-

ence of ambush predators, tadpoles become acceleration/

maneuver specialists, while in the presence of run-down

predators, tadpoles become speed specialists. Morpho-

logical adaptations for such escape pathways include a

broader tail (Lardner 1998; Laurila et al. 2006; Relyea

2002; Relyea 2003; Sosa et al. 2009; Teplitsky et al.

2003) or a longer tail, respectively (Higginson and Rux-

ton 2010; Moore et al. 2004; Relyea 2000). In some cas-

es, the presence of predators causes early metamorpho-

sis (Benard 2004; Higginson and Ruxton 2010; Relyea

2007; Werner 1986).

Morphological changes in response to predator pres-

ence occur in a diversity of amphibian taxa that are dis-

parate both in phylogenetic and life-history traits. Frog

species possessing different life-history traits show dif-

ferent anti-predator responses to different predators and

competitors (Laurila et al. 2006; Relyea 2001a; Relyea

2001b; Relyea and Yurewicz 2002). For fully aquatic

tadpoles, these morphological responses are now well

known.

Laying eggs away from water in a foamy mass, in

which tadpoles develop up to a pre-metamorphic stage

before falling into water, is an alternative life history strat-

egy, often known as foam nesting (Duellman and Trueb

1986). This strategy is considered to facilitate predator

avoidance of eggs and early-stage tadpoles (Hodl 1992;

Magnusson and Hero 1991), and to reduce the duration

of the larval stage (through rapid development during the

out-of- water phase).

The Hourglass treefrog {Polypedates cruciger), a Sri

Lankan endemic, shows a derived reproductive strategy

from aquatic egg deposition. These frogs make foamy

nests overhanging water bodies, in which they lay their

eggs. Tadpoles develop within the nest, up to Gosner

stage 23 and then fall into water, where they undergo

further development reaching metamorphosis. Adult P.

cruciger are arboreal, but sometimes visit pools at night,

apparently to rehydrate.

Correspondence. Email: ^madhava_m@ mac.com

amphibian-reptile-conservation.org

014

November 2011 I Volume 5 I Number 2 I e29

Ariyasiri et al.

Figure 1. Outline of tadpole (lateral and dorsal views), depicting measurements that were used in this study; total length (TL), tail

length (TAL), maximum tail height (MTH), maximum tail height to tip of tail (MTH-t), total muscle height (TMH), total muscle

width (TMW), body length (BL), inter-orbital distance (lOD), internasal distance (IND), and limb length (LL).

Fish prey on such early-stage tadpoles that fall into

water (this has been documented for other species, in

which tadpoles of arboreal gel-encapsulated egg layers

fall into water and are eaten by various aquatic preda-

tors; Magnusson and Hero 1991). Tadpoles of P. cruci-

ger are preyed on by various fish species, including the

Combtail, Belontia signata (Belontiidae), the Snake-

head, Channa orientalis (Channidae), and nonnative

and introduced Guppy, Poecilia reticulata (Poeciliidae;

M. Meegaskumbura, pers. obs.). This study tests the de-

velopmental response of P. cruciger tadpoles to aquatic

predation pressure.

Methods and materials

A single foam nest of Polypedates cruciger attached to a

twig above a pond was observed in Peradeniya Univer-

sity Gardens, Sri Lanka (7°15’34.02”N, 80°35’49.71”E;

600 m asl). Tadpoles that emerged six days after the foam

nest was first made (fertilization was observed) were

reared in a glass aquarium for seven days, until the ex-

periment began.

The experimental setup was as follows: eleven

equally sized glass aquaria (size: 45 x 30 x 30 cm) each

with 25 tadpoles was set up. Three of these were used as

controls, and contained only tadpoles. Of the eight ex-

perimental aquaria, four contained tadpoles and fish, but

visual contact between the tadpoles and fish was prevent-

ed by an opaque, water-permeable screen so that they

shared the same water (chemicals produced by fish or

tadpoles could thus be detected by any individual in the

aquarium); these treatments were termed “closed” (they

were established to provide tadpoles with an attenuated

predator presence). The other four aquaria contained

both tadpoles and fish, but allowing for visual (though

not physical) contact between the predators and potential

prey. They too, shared the same water, and were termed

“open.”

All other experimental conditions were kept identi-

cal for all tanks. The fish and tadpoles were fed a pro-

tein-rich aquarium-fish food. Daily partial water changes

were made using water from an animal-free aquarium

that had a UV-C sterilizer (to remove pathogenic organ-

isms) and an aerating power filter (to aerate water and

remove traces of chlorine and ammonia that could be

present in tap water).

Samples were taken 12 days after the beginning of

the experiment. They were anesthetized in MS222 and

measured using a vernier caliper under a stereo micro-

scope. Six tadpoles were sampled arbitrarily from each

replicate. They were measured to +0.01 mm using a digi-

tal caliper. The following measurements were taken: to-

tal length (TL), tail length (TAL), maximum tail height

(MTH), maximum tail height to tip of tail (MTH-t), total

muscle height (TMH), total muscle width (TMW), body

length (BL), inter-orbital width (lOD), and intemasal

distance (IND; Fig. 1).

amphibian-reptile-conservation.org

015

November 2011 I Volume 5 I Number 2 I e29

Plasticity in tadpoles of Polypedates cruciger

Figure 2. The morphology of early tadpole stages: A, control; B, “open.” Scale bar 1 mm.

Coefficients of variation (CV scores) were deter-

mined and variables that had CV > 5%, and individuals

that were outliers, were excluded from analyses. Prior

to all analyses (except determination of CV scores) data

were normalized through log^^ transformation. The mean

of each replicate was used in the subsequent analyses.

Systat version 11.00.01 for Windows XP was used

for the statistical analysis. Principal Components Analy-

sis (PCA) of means of character covariance matrix was

used to reduce the dimensionality of morphological

variables and to identify variables that may discriminate

between the treatments. Different axis rotations were

tested, and the one that yielded optimal interpretability

of variation among variables is reported.

Discriminant Functions Analysis (DFA) was carried

out to distinguish between the three experimental groups.

To visualize relationships between the variables of

tadpole morphology, box plots depicting mean and stan-

dard error were made.

Results

Variables having CV scores > 5%, IND and LL, were ex-

cluded, leaving seven variables (TL, TAL, MTH, MTH-t,

TMH, TMW, and BL) available for further analysis.

In the PC space of unrotated PC 1 and PC2 axes,

the two treatments (“closed” and “open”), and the “con-

PC 1

Figure 3. Principal components space of PCI vs. PC2 (un-

regressed) of tadpole measurements in the two experimental

conditions (“open” and “closed”) and the controls of the early

sampling regime. The PCI axis, which explains 46% of the

variance, is mostly represented by tail length, total length, and

inter-orbital width. The PC2 axis, which explains 24% of the

variance, mostly represents tail height-related variables.

amphibian-reptile-conservation.org

016

November 2011 I Volume 5 I Number 2 I e29

Ariyasiri et al.

Figure 4. Canonical variables plot of discriminant function

analysis (unregressed) of the two experimental conditions

(“open” and “closed”) and the control. Ninety -five percent con-

fidence elipses of these three do not overlap with one another,

and are centered on the centroid each group.

trol” tadpoles separate well (Figs. 3, 4). On the PC 1

axis, which explains 46% of the variance, several vari-

ables representative of tail and total lengths, and lOD

load heavily (component loadings: TAL = 0.889, MTH-t

= 0.871, lOD = 0.869, TL = 0.825; TMW = 0.667; Table

1). On this axis, “control” and “open” do not overlap,

but “closed” overlaps with both the former cases and is

placed in between these. Hence, presence of fish seems

to increase total and tail-length related dimensions in

tadpoles. On the PC 2 axis, which explains 24% of the

variance, “closed” does not overlap with either “open” or

“control.” However, both “open” and “control” overlap

with each other completely on this axis, which is mostly

explained by tail height-related variables (component

loadings TMH = 0.811, MTH = 0.624; Table 1). Con-

sidering unrotated PC 1 vs. PC 3, PC 1 vs. PC 4, PC 2

vs. PC 3, and PC 2 vs. PC 4 for these, the treatments and

controls overlap with each other to various degrees on

the PC 3 and PC 4 axes (not shown) but, as explained

above, not on the PC 1 and PC 2 axes.

The Discriminant Functions Analysis shows that the

95% confidence ellipses do not overlap with each other

(Fig. 4).

Some of the tail-length associated variables (means

and standard errors) (TAL, MTH-t, TL, and TMW) show

distinctions among the three groups; only the box plot of

MTH-t is shown (Fig. 5).

Discussion

Because of predation, developmental anomalies, patho-

gens, and unfavorable environmental conditions, not all

amphibian larvae develop to metamorphosis. Often en-

tire egg clutches are destroyed even before tadpoles be-

come free swimming.

Predation reduction of egg and early stage tadpoles

has been suggested to have driven the evolution of egg

deposition out of water for many forms (Doughty 2002).

This hypothesis is plausible, but predator avoidance is

still important even after early-stage tadpoles of foam-

nesting species fall into water. Indeed, we have observed

tadpoles of P. cruciger being preyed upon by various fish

species. Once a falling tadpole is detected by predatory

fish, it lurks under the nest waiting for more tadpoles

to fall (M. Meegaskumbura, pers. obs.). In such a situ-

ation, there is clearly an advantage for tadpole’s ability

to evacuate the impact area as soon as possible. We have

observed this: tadpoles of P. cruciger, upon impacting

the surface of the water, quickly react by swimming

away rapidly, in an apparently arbitrary direction, until

Table 1. Component loadings for axes 1 -4 for the Principal Component Analysis, variance explained, and percentage of total vari-

ance explained for early sample treatments and controls (unregressed: “open,” “closed,” and “control”).

Component Loadings

TAL

0.889

-0.341

-0.160

0.241

MTHT

0.871

-0.188

-0.297

-0.232

lOD

0.869

0.332

0.007

-0.312

0.825

-0.466

0.112

0.281

TMW

0.667

0.477

0.349

0.374

TMH

-0.102

0.811

-0.242

0.464

MTH

0.407

0.624

0.514

-0.370

-0.188

-0.401

0.874

0.128

Variance Explained by Components

3.642

1.914

1.335

0.796

% of Total Variance Explained

45.530

23.929

16.686

9.955

amphibian-reptile-conservation.org

017

November 2011 I Volume 5 I Number 2 I e29

Plasticity in tadpoles of Polypedates cruciger

Figure 5. Boxplot depicting the means and standard errors of

the two treatments (“open” and “closed”) and the control.

they reach a safe submerged refuge. Furthermore, even

though young tadpoles are attached by their cement

glands to underwater substrates at this stage, they react

quickly to any disturbance by fast and apparently ran-

dom swimming (M. Meegaskumbura, pers. obs.). These

observations are indications that effective swimming is

an important survival attribute in tadpoles.

PCA and DFA results are complementary and show

tadpoles of the “control” and “open groups” to be diver-

gent in body morphology. It is known that a larger body

confers reduced risk of predation (Buskirk and Schmidt

2000), as this enables animals to swim faster, or acceler-

ate and maneuver better. The “open” body morphology

of P. cruciger tadpoles matches the features of tadpoles

from other unrelated taxa that respond to predation by

achieving a fast-swimming body morphology e.g., lon-

ger tail, greater total length: Buskirk and Relyea (1998);

Teplitsky et al. (2003).

Behavioral plasticity might be inexpensive due to

absence of a need for new or altered structures to meet

new challenges (Buskirk 2002). Though behavioral re-

sponse of tadpoles to predators was not quantified in

this study, we observed that tadpoles from “open” tanks

reacted most swiftly to disturbances when compared to

“closed” and “control” groups.

We have yet to study the effects of predator presence

on early metamorphosis, something that several other

authors have previously reported on (Gomez-Mestre et

al. 2008; Lardner 1998; Vonesh and Warkentin 2006). If

early metamorphosis occurs in tadpoles that develop in

association with a predator, the resulting tadpoles may

have a smaller body (Lardner 1998).

Although our data demonstrate that P. cruciger tad-

poles exhibit predator-induced plasticity, they reveal little

about the patterns of plasticity. For example, we do not

know whether all tadpole stages show predator induced

plasticity, or if the presence of predators induces early

metamorphosis. Further experimentation is warranted.

Multiple layers of protection, initially through har-

boring of the vulnerable early developmental forms in

a foam nest, and later, after partially developed tadpoles

amphibian-reptile-conservation.org

fall into water, in the accelerated development responses

to aquatic predator presence, seem like adaptations to

help survive in a predator high environment. If foam nest-

ing evolved as a response to predator avoidance of early

tadpole stages, it can be argued that there was a heavy

predation cost for the aquatic larvae, at least historically.

Then even the partially developed tadpoles would have

to face some form of predation, from the very predators

that would have eaten them as early-stage larvae, had the

eggs been laid in water, even though at a reduced inten-

sity. These adaptations could be a reason for the wide

distribution of this species across several habitat types in

the wet and the intermediate zone of Sri Lanka. It will be

interesting to determine whether adaptations observed in

P. cruciger are seen also in tadpoles of Taruga, its sister

genus (Meegaskumbura et al. 2010).

Introduced predatory fishes may have various feed-

ing mechanisms, which tadpoles living in these waters

may not be adapted to. For instance, to avoid predation

from an ambush predator, an accelerating or maneuver-

ing tadpole body form may be needed. If this is not pres-

ent, an introduced form may destroy whole populations

of tadpoles.

Hence, when causes for decline of amphibians are

considered in the context of to introductory predatory

fishes or aquatic predators, study of tadpole morphologi-

cal adaptability may be important to determine the actual

mechanisms of decline.

Acknowledgments. — We wish to thank the two anon-

ymous reviewers for their valuable comments improving

the paper. We acknowledge the Department of Wildlife

Conservation of Sri Lanka for research permits to study

tadpoles, and the Department of Zoology, Faculty of

Science, University of Peradeniya, for resources. We

are grateful to the Amphibian Specialist Group (lUCN/

SSC), Global Wildlife Conservation (GWC), Amphibian

Redlisting Authority (ARLA/IUCN/SSC), Rohan Pethi-

yagoda, Kelum Manamendra-Arachchi, Don Church,

and James Lewis for supporting this work.

Literature cited

Anderson AL, Brown WD. 2009. Plasticity of hatching in

green frogs (Rana clamitans) to both egg and tadpole preda-

tors. Herpetologica 65(2);207-213.

Benard MF. 2004. Predator-induced phenotypic plasticity in

organisms with complex life histories. Annual Review of

Ecology, Evolution, and Systeniatics 35:651-673.

Buskirk JV. 2002. Phenotypic lability and the evolution of pred-

ator-induced plasticity in tadpoles. Evolution 56(2) ; 361-

370.

Buskirk JV, Relyea RA. 1998. Selection for phenotypic plas-

ticity in Rana sylvatica tadpoles. Biological Journal of the

Linnean Society 65(3);301-328.

018

November 2011 I Volume 5 I Number 2 I e29

Ariyasiri et al.

Buskirk JV, Schmidt BR. 2000. Predator-induced phenotypic

plasticity in larval newts: trade-offs, selection, and variation

in nature. Ecology 81(ll):3009-3028.

Doughty P. 2002. Coevolution of developmental plasticity

and large egg size in Crinia georgiana tadpoles. Copeia

2002(4):928-937.

Duellman we. Truer L. 1986. Biology of Amphibians. Mc-

Graw-Hill, New York. 696 p.

Gomez-Mestre I, Wiens JJ, Warkentin KM. 2008. Evolu-

tion of adaptive plasticity: risk-sensitive hatching in neo-

tropical leaf-breeding tree frogs. Ecological Monographs

78(2):205-224.

Higginson ad, Ruxton GD. 2010. Adaptive changes in size

and age at metamorphosis can qualitatively vary with preda-

tor type and available defenses. Ecology 91(9):2756-2768.

Hodl W. 1992. Reproductive behavior in the neotrpical foam-

nesting frog Pleurodema diplolistris (Leptodactylidae).

Amphibia-Reptilia 13(3): 263 -274.

Eardner B. 1998. Plasticity or fixed adaptive traits? Strategies

for predation avoidance in Rana arvalis tadpoles. Oecologia

117(1-2):119-126.

Eaurila a, Pakkasmaa S, Merila J. 2006. Population diver-

gence in growth rate and antipredator defences in Rana ar-

valis. Oecologia 147(4):585-595.

Magnusson we. Hero J-M. 1991 . Predation and the evolution

of complex oviposition behaviour in Amazon rainforest

frogs. Oecologia 86(3):310-318.

Meegaskumbura M, Meegaskumbura S, Bo watte G, Mana-

mendra-Arachchchi K, Pethiyagoda R., Hanken J, Sch-

neider CJ. 2010. Taruga (Anura: Rhacophoridae), a new ge-

nus of foam-nesting tree frogs endemic to Sri Lanka. Ceylon

Journal of Science (Biological Sciences) 39(2):75-94.

Moore RD, Grieeiths RA, O’Brien CM, Murphy A, Jay

D. 2004. Induced defences in an endangered amphibian

in response to an introduced snake predator. Oecologia

141(1):139-147.

Relyea RA. 2000. Trait-mediated indirect effects in larval an-

urans: reversing competition with the threat of predation.

Ecology 81(8): 2278-2289.

Relyea RA. 2001a. Morphological and behavioral plasticity of

larval anurans in response to different predators. Ecology

82(2):523-540.

Relyea RA. 2001b. The relationship between predation risk

and antipredator responses in larval anurans. Ecology

82(2):541-554.

Relyea RA. 2002. Costs of phenotypic plasticity. The American

Naturalist 159(3):272-282.

Relyea RA. 2003. Predators come and predators go: the revers-

ibility of predator-induced traits. Ecology 84(7): 1840-1848.

Relyea RA. 2007. Getting out alive: how predators affect the

decision to metamorphose. Oecologia 152(3):389-400.

Relyea RA, Yurewicz KL. 2002. Predicting community out-

comes from pairwise interactions: integrating density- and

trait-mediated effects. Oecologia 131(4):569-579.

Sosa JA, Ryan MJ, Schlaepeer MA. 2009. Induced morpho-

logical plasticity in Lowland leopard frog larvae (Rana

yavapaiensis) does not confer a survival advantage against

Green sunfish (Lepomis cyanellus). Journal of Herpetology

43(3):460-468.

TeplitskyC,Plenet S, JolyP. 2003. Tadpoles’ responses to risk

offish introduction. Oecologia 134(2):270-277.

VoNESH JR, Warkentin KM. 2006. Opposite shifts in size at

metamorphosis in response to larval and metamorph preda-

tors. Ecology 87(3): 556-562.

Werner EE. 1986. Amphibian metamorphosis: growth rate,

predation risk, and the optimal size at transformation. The

American Naturalist 128(3):3 19-341.

Manuscript received: 13 April 2010

Accepted: 25 October 2011

Published: 12 November 2011

KRISHAN ARIYASIRI graduated from the University of

Peradeniya in 2008 where he studied the vertebrate diversity

changing with elevation gradient along the Maha-Oya, Hantana

forest during his senior year. His diverse interests in biology

range from ecology, Raptor biology, microhabitat associations

of frogs, and morphological adaptability in amphibians. He is

currently contemplating graduate studies in molecular genetics.

amphibian-reptile-conservation.org 019 November 2011 | Volume 5 | Number 2 | e29

Plasticity in tadpoles of Polypedates cruciger

GAYAN BOWATTE graduated from the University of Perad-

eniya in 2009. Gayan is currently a graduate student at the Post-

graduate Institute of Science (Peradeniya) and works on nitro-

gen-based stressors affecting amphibians. His interests include

systematics and morphophological development of tadpoles.

SUYAMA MEEGASKUMBURA is a Senior Eecturer at the

Department of Zoology, Faculty of Science, University of Per-

adeniya. She is an evolutionary biologist, mammalian biologist,

and parasitologist. Suyama was awarded the B.Sc. in Zoology

(with first class honours), M.Sc. in Parasitology (University of

Peradeniya), and a Ph.D. in Biology from Boston University.

Her research over the past decade has been on molecular sys-

tematics, evolutionary biology, and ecology of small mammals

and parasites. She has described a new species of shrew from

the Sinharaja World Heritage Site and Momingside, and has re-

vised the taxonomy of several other small mammal taxa, mostly

using molecular systematics. She is the sub-editor of the Cey-

lon Journal of Science, a journal that publishes peer-reviewed

research work of South Asian biologists. She sits on various

education boards that are concerned with graduate student edu-

cation at the University of Peradeniya and the Postgraduate In-

stitute of Science.

UDENI MENIKE graduated from the University of Peradeniya

in 2008. She studied the species composition and prevalence of

external parasites of Suncus murinus (Soricidea: Crocidurinae)

on the University of Peradeniya premises, for her final year re-

search project. Currently she is working on developing non-

destructive sampling methods for small mammals.

amphibian-reptile-conservation.org

020

November 2011 I Volume 5 I Number 2 I e29

Ariyasiri et al.

MADHAVA MEEGASKUMBUP^ is currently a Senior Eec-

turer at the Department of Zoology, Faculty of Science, Uni-

versity of Peradeniya, Sri Eanka. He is an evolutionary biolo-

gist and ecologist by training and received his B.Sc. in Zoology

from the University of Peradeniya and a Ph.D. from Boston

University (2007). Upon receiving his doctorate degree he was

a Ziff Environmental Postdoctoral Fellow for two years at Har-

vard University (Harvard University Center for the Environ-

ment and Museum of Comparative Zoology). Over the past

decade he has done research on systematics and phylogenet-

ics, evolution, and ecology of Sri Eanka’s frogs, mammals, and

fish. Madhava is the Co-Chairman of the Amphibian Specialist

Group Sri Eanka (ASGSE/IUCN/SSC) and a member of the

Amphibian Redlisting Authority (AREA/IUCN/SSC). He has

published about 20 peer-reviewed papers, several book chap-

ters, and popular articles. He has described about 20 new spe-

cies of Sri Eankan animals (frogs, fish, and a mammal) and a

new frog genus (Taruga).

amphibian-reptile-conservation.org

021

November 2011 I Volume 5 I Number 2 I e29

the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any

medium, provided the original author and source are credited.

Amphibian & Reptiie Conservation 5(2):22-32.

Morphology and ecology of Microhyla rubra (Anura:

Microhylidae) tadpoles from Sri Lanka

^GAYAN BOWATTE AND ' ^MADHAVA MEEGASKUMBURA

^Department of Zoology, Faculty of Science, University of Peradeniya, SRI LANKA

Abstract . — ^The life-history, ecology, external and buccal morphology of Microhyia rubra (Jerdon,

1854) tadpoles are described. Approximately 400 eggs, ready to hatch, were observed as a single

mass and several of these were reared in laboratory. Tadpoles showed several characters that are

not seen in most other microhylids: a whip-like tail-end flagellum, a dorsoterminal mouth, a trans-

parent body, absence of flaps and existence of a median notch on upper lip, presence of papillae (or

scallops) on lower lip, and a deep ventral tail fin (compared to the dorsal tail fin). Microhyia rubra

tadpoles also have several features, so far not noted in other microhylids: six papillae (or scallops)

on lower oral flap, a crescent-shaped spiracular opening, and an enlarged crest on ventral tail fin.

For some characters, such as shape of the oral flaps, we show that there is considerable varia-

tion within and between Gosner stages. This species deposits its eggs as rafts in ephemeral pools

where water chemistry (bound ammonia, salinity, conductivity, pH, sulphate ion concentration) and

temperature are apparently favorable for rapid growth, reducing the risk of predation from fully

aquatic predators. Since oxygen concentrations in these habitats are low and free ammonia concen-

trations are moderately high, occupying surface layers of pools would enable the eggs and tadpoles

to overcome these impediments to growth and survival.

Key words. Microhylinae, microhyline tadpoles, morphology, buccal, ecology, Microhyia rubra

Citation: Bowatte G, Meegaskumbura M. 2011 . Morphology and ecology of Microhyia rubra (Anura: Microhylidae) tadpoles from Sri Lanka. Amphibian &

Reptile Conservation 5(2):22-32(e30).

Introduction

The natural history and reproductive biology of microhy-

lid frogs are poorly known (Wassersug 1980; Donnelly

et al. 1990; Lehr et al. 2007). Although descriptions of

tadpole characters useful in taxonomy have been de-

scribed only for a few species, tadpole morphology var-

ies considerably both inter- and intra-specifically (Don-

nelly et al. 1990). Hence, it is important to study tadpole

morphology in greater detail, making inter-species com-

parisons more useful for phylogenetic and comparative-

morphological analyses.

The Red narrow-mouthed frog, Microhyia rubra, is

widely distributed in the lower elevation regions of Sri

Lanka, peninsular India, and Bangladesh, rarely occur-

ring above 500 m asl (Kirtisinghe 1957; Manamendra-

Arachchi and Pethiyagoda 2006; lUCN 2004); it is found

predominantly in drier parts of these countries. The spe-

cies is often found under logs, piles of rubble, haystacks,

and stones, where comparatively higher moisture levels

exist. Small size, nocturnal habits, and cryptic nature of

these frogs make them difficult to encounter in the field.

Nonetheless, Microhyia rubra is categorized as

“Least Concern” by the lUCN, due to its wide distribu-

Correspondence. Email: ^madhava_m@ mac.com

tion, tolerance of dry environmental conditions, and high

population densities.

Despite their abundance, details of the life history of

Microhyia rubra, especially tadpole characteristics and

biology, are still poorly known. Several previous workers

(Rao 1918; Parker 1928, 1934; Kirtisinghe 1957, 1958)

have described the external morphology of the tadpoles,

and Rao (1918) states that they are not transparent. Kir-

tisinghe, (1957) provided a brief description of the ex-

ternal morphology of the tadpole, including presence of

a tail-end fiagellum, dorso-terminal mouth, spiracular

opening above a notched fiap on underside of the belly,

and the deep lower crest of the ventral tail fin. Kirtisinghe

(1957) provides a drawing of oral fiaps, but without a

description. Internal buccal morphology is not discussed

by any of these researchers.

Here we provide a more complete description of the

external morphology of Microhyia rubra tadpoles and

provide the first description of their buccal morphology.

We particularly concentrate on the mouth location, spir-

acle location, shape of spiracular opening, tail morphol-

ogy, and mouthparts, as these features are shown to vary

considerably among and within microhylids (Donnelly et

al. 1990) and are of potential importance in systematics.

amphibian-reptile-conservation.org

022

December 2011 I Volume 5 I Number 2 I e30

Bowatte and Meegaskumbura

Figure 1. Open and shallow ephemeral pool lined by grass and shrubs, where floating eggs were sampled.

Methods and materials

Location (08°16’49.43” N, 80°28’49.96” E): Several

eggs in late embryonic stages were collected (identity

of species was not known at time of collection) from an

ephemeral man-made pool near Nachchaduwa reservoir

in Anuradhapura (Fig. 1). Tadpoles at Stage 24 (Gosner

1960) emerged from these eggs after two days. These

tadpoles were raised in the laboratory, with partial daily

water changes of dechlorinated water, and periodically

sampled until metamorphosis. Tadpoles were fed on

boiled egg yolk. Metamorphs were raised an additional

month, and identified using taxonomic keys devised for

adult frogs (Manamendra-Arachchi and Pethiyagoda

2006). Tadpoles were fixed in 10% buffered formalin for

two days and preserved in a 1 : 1 mixture of 10% buffered

formalin and 70% alcohol. Tadpoles are deposited in the

collection of the Department of Zoology, University of

Peradeniya, Sri Lanka (DZ).

Grillitsch et al. (1993) and McDiarmid and Altig

(1999) were followed for external description of tad-

poles. For internal oral anatomy, a combination of Khan

(2000) and Wassersug (1976) was followed. The surgical

method delineated by Wassersug (1976) was used and

the following measurements were taken (Fig. 2): maxi-

mum height of body (bh), maximum width of body (bw),

maximum diameter of eye (ed), maximum height of tail

(ht), maximum height of lower tail fin (If), internarial dis-

tance (nn), naro-pupilar distance (np), interpupilar dis-

tance (pp), rostro-narial distance (rn), distance from tip

of snout to opening of spiracle (ss), distance from tip of

snout to insertion of upper tail fin (su), snout-vent length

(svl), total length (tl), maximum height of upper tail fin

(uf), distance from vent to tip of tail (vt), tail muscle

height (tmh), and tail muscle width (tmw). Morphol-

ogy was observed using a Mode zoom-stereomicroscope

(6-50 x). Tadpoles were measured using digital calipers

(measured to the nearest 0.01 mm).

Results

Description of tadpole

External morphology. The following description is based

on five Stage 35 tadpoles of Microhyla rubra (DZ 1033-

37) except where explicitly stated.

In dorsal view, body clearly differentiated into two

parts, a longer and wider anterior region (Rl) and a nar-

rower posterior region (R2). Anterior region almost twice

as long and wide as posterior region (Figs. 2 and 3). Eyes

amphibian-reptile-conservation.org

023

December 2011 I Volume 5 I Number 2 I e30

Morphology and ecology of tadpoles, Microhyla rubra

Figure 2. Outline of Microhyla rubra tadpoles showing the measurements that were taken.

small (ed/bw = 0.22) and snout rounded. Head and body

posterior to eyes with sides parallel to each other, and

conjunction of R1 and R2 forms an angle of 137-148°.

Eyes directed slightly dorsolaterally, bulbous, and entire

eye visible through epidermis due to dearth of pigmen-

tation. Nares closed (nn/pp = 0.21), narial depressions

visible, oval, unpigmented to slightly pigmented, located

immediately anterior to two small concentrated patches

of pigment, anterodorsolaterally directed, and closer to

snout tip than to pupils. Nasolacrimal duct apparent. A

lateral protruding ridge anterior to eye. Mouth narrow,

superior, lower and upper-lips both visible. Tail long, ta-

pering, with a whip-like flagellum (pointed tail tip; Fig.

4).

In profile, R1 wedge-shaped, pointed at snout, an-

terior-dorsal aspect straight, and anterior-ventral aspect

slightly rounded. R2 ventrally rounded and dorsally

slightly rounded. Gut contained in R2, overlaid with iri-

dophores (Fig. 3E). A paired gas-fllled cavities present

dorsolateral to the gut (probably the developing lungs);

horizontal dark bar located dorsal to gas-fllled cavities.

Spiracle mid-ventral, transparent, ends at posterior ven-

tral part of body, dorsally attached to body wall, and

ventrally free with a small posteriorly extending flap

with medial notch near vent. Ventral tail fin begins at the

dorsal attached end of the spiracular opening. Spiracular

opening crescent-shaped with anterior portion of the ven-

tral tail fin contained within the spiracle (Fig. 3C). Vent

tube in lower tail fin, posterior to spiracle opening. Tail

musculature weak, extending to end of tail tip (tail-mus-

cle height/body height = 0.43; tail-muscle width/body

width = 0.31), V-shaped myomeres apparent only in pos-

terior two-thirds of tail (Fig. 3A). Dorsal tail fin deeper

than ventral tail fin, both fins originate above and below

the same vertical point on body. Fins reduced towards

end, proximally a deep convex extension of ventral tail

fin (lowest crest) distally, a smaller crest towards middle

of tail (Fig. 5).

In ventral view, eyes barely visible, but silhouette of

eye-ball apparent through unpigmented skin. Extended

flap of lower lip visible. Coiled gut visible, positioned

slightly to left of midline, overlaid with iridophores.

Heart at boundary of R1 and R2.

Oral flaps: upper lip not fleshy (Fig. 3B), with a

slight medial notch. Edge of lower lip slightly scalloped,

with three projections on each lobe (Fig. 8).

Buccal morphology

Eabial keratinized teeth were absent in all individuals

examined.

Ventral buccal region. Prelingual arena U-shaped,

length greater than width, curved portion of U directed

anteriorly toward oral aperture. A pair of dorsally-direct-

amphibian-reptile-conservation.org

024

December 2011 I Volume 5 I Number 2 I e30

Bowatte and Meegaskumbura

Figure 3. Microhyla rubra tadpole (Stage 38) in life showing: (A) the long tail with a distinct flagellum, (B) position of mouth, (C)

shape of the spiracle and position of the vent tube in tail, (D) Shape of the convex curvature in ventral fin, and (E) close up of the

head and body showing the nasolacrimal duct, distribution of pigmentation, mouth position, and groove on non-fleshy upper lip.

amphibian-reptile-conservation.org

025

December 2011 I Volume 5 I Number 2 I e30

Morphology and ecology of tadpoles, Microhyla rubra

Figure 4. Dorsal aspect of the body and part of the tail of a Microhyla rubra tadpole (Stage 35). Scale bar, 1 mm.

ed lateral infralabial papillae of equal size line mouth

opening. Fleshy fold on the lateral walls of mouth open-

ing. A fleshy fold on mouth floor posterior to infralabial

papillae, directed towards buccal cavity. A pair of lat-

eral buccal pockets in anterior region of buccal floor. A

single pair of small papillae on anterior wall of buccal

cavity, on either side of mouth aperture, not attached to

tongue. Conical, non-papillated tongue anlage, broader

anteriorly, without pigment, narrower and free posteri-

orly, with pigment. Buccal floor arena (BFA) triangular,

laterally elevated, medially depressed, forming a narrow

passage at the anterior portion of BFA, posterior end of

buccal floor much broader than anterior end. Two small

and blunt, two large, and one medium-sized symmetrical

pairs of conical BFA papillae. Small papillae (length =

0.07 mm) anterior to all others. Medium papillae (length

= 0.16-0.19 mm) close to glottis. Large papillae (length

= 0.27-0.34 mm) further from glottis, posterior to me-

dium papillae. Single conical large medial preglottal pa-

pilla. Buccal pockets long and narrow, sickle- shaped, and

blunt at the blind end. A pair of symmetrical, small blunt

proximal prepocket papillae. Pairs of one large conical,

three medium conical, four small blunt postpocket papil-

lae. A large conical medially curved distal and sinistral

prepocket papilla. A large and medium conical, medi-

ally curved, distal dextral prepocket papilla. Trachaea

club-shaped, protruding from base of velum, extending

to base of BFA, ending in elevated lips. Broad ventral

velum without strong spicular support, free margin of ve-

lum smooth, covered by secretory pits, and containing a

single broad projection above third Alter plate (Fig. 6).

Dorsal buccal region. Choanae blind ended. Pre-

narial arena a posteriorly-directed V-shaped depression.

Prenarial papilla, single, medial, small, blunt, placed an-

terior to narial papilla. Narial papillae hang from narial

depression, slightly twisted, long, flat, robust, with three

projections towards the anteriorly-directed tip; the mid-

dle projection longest. Postnarial ridge slightly serrated.

Buccal roof arena (BRA) triangular, broad anteriorly, and

lined by postero-lateral BRA border with papillae. Close

Table 1. Means and standard deviations of 12 tadpole body measurements of M. rubra at different Gosner stages (26, 31, 33,

and 35).

Characteristics

Stage 26

Stage 31

Stage 33

Stage 35

n = 2

n = 6

Body height (bh)

2.45 + 0.02

3.63 + 0.01

4.60 + 0.15

5.54 + 0.67

Body width (bw)

2.83 + 0.37

4.47 + 0.06

5.79 + 0.21

6.41 + 0.66

Maximum taii height (th)

2.98 + 0.32

4.49 + 0.32

5.24 + 0.04

6.26 + 1.07

Inter narial distance (nn)

0.64 + 0.01

0.89 + 0.01

1.07 + 0.02

1.23 + 0.12

Inter popular distance (pp)

2.68 + 0.37

4.20 + 0.09

5.50 + 0.22

5.94 + 0.83

Snout-vent length (svi)

4.24 + 0.09

5.85 + 0.30

7.40 + 0.34

8.67 + 1.22

Total length (tl)

14.48 + 1.65

20.59 + 2.47

26.23 + 0.55

29.00 + 3.11

Vent to tail tip length (vt)

10 . 24 + 1.75

14.74 + 2.18

18.83 + 0.21

20.39 + 2.01

Tail muscle height (tmh)

1.07 + 0.13

1.93 + 0.30

2.23 + 0.01

2.35 + 0.24

Tail muscle width (tmw)

0.66 + 0.01

1.33 + 0.09

1.69 + 0.24

1.98 + 0.29

amphibian-reptile-conservation.org

026

December 2011 | Volume 5

Number 2 | e30

Bowatte and Meegaskumbura

Figure 5. Profile of the whole body of the Microhyla rubra tadpole (Stage 35). Scale bar, 1 mm.

to BRA apex, one pair long (length = 0.44-0.47 mm)

and pointed; one pair medium (length = 0.14-0.19 mm)

and pointed; BRA papillae, lateral to apex; BRA border

with a few small (length = 0.04-0.06 mm) BRA papillae.

Broad roof glandular area anterior to dorsal velum and

dorsal velum gradually thins medially (Fig 7).

Ventral pharynx region. Branchial baskets triangu-

lar, half of the filter cavities anterior to the velum, and

all three filter plates distinct. A distinctly ridged oval to-

rus present in each filter cavity and subvelar surface with

many secretory ridges (Fig. 6).

Color in life. Body transparent and light yellowish

grey. In profile, dorsum densely pigmented compared to

venter, pink region present between eyes and coiled gut.

Iris silver, with dark inverted V- shape at ventral edge. R2

studded with silver iridopores and dark-brown pigment

cells (Fig. 3E). Tail fins lightly pigmented in dark brown.

Figure 6. Ventral buccal morphology of the Microhyla rubra

tadpole (Stage 35). Scale bar, 1 mm.

Tail musculature equally pigmented throughout, size of

pigment patches reducing posteriorly (Fig. 3A, B, C, and

D). Upper margin of the hind limb and toes pigmented

(Fig. 3A, C, and D). In dorsal view, densely pigmented

areas located near nasal openings, between nasal opening

and point of origin of upper tail fin, along the base of the

upper tail fin and in the gas-filled cavities. Posterior to

nasal markings a red band extends to margin of R1 and

R2. Eyeballs apparent and black in color.

Color (preserved). Body semi-transparent to brown-

ish-white, tail lighter color than the body. Pigments on

body star-shaped, giving the appearance of powder coat-

ing. Higher densities of pigments occur dorsally than

ventrally. A median symmetrical dorsal band of dark

brown to black melanophores covers the brain region

and extend to near the base of eyes and nasal pits. Dark

brown to black pigment patches present posteriorly to

low-pigmented nasal depressions. Iris silver, with scat-

tered dark patches. Two narrow dark lines originate at

dorsal pole of pupil and extend ventrally. Symmetrical

black bands over dorsum to gas-filled cavities at the ori-

gin of the tail musculature. A dark brown line runs along

the top of the tail musculature between dark bands of

gas-filled cavities. R2 (Fig. 2) in the body almost covered

with iridiophores, giving it a characteristic silvery shine,

and black color patches present on this silver region. Re-

duced pigmentation in the tail musculature and tail fins.

Ventrally, heart visible, cream colored, at margin of R1

and R2.

Variation. There is a substantial amount of variation

in the lower lip in tadpoles of different developmental

stages, and sometimes even within a given developmen-

tal stage. At Stage 25 (early stage) for instance, there is a

single pair of scallops on the lower lip but these develop

into six very distinct papillae (three pairs) by late Stage

25. At Stage 30, the scallops are distinct and there is little

variation within the stage. By Stage 35, the scallops are

not clearly discernible (and there is little variation within

the stage; Fig. 8). The tail-fin shape changes from a sim-

ple long triangular shape (Stage 25) to a more complex

shape with two crests on the ventral tail fin (anterior crest

deeper and crest in middle of tail shallower; Stage 35).

amphibian-reptile-conservation.org

027

December 2011 I Volume 5 I Number 2 I e30

Morphology and ecology of tadpoles, Microhyla rubra

Figure 7. Dorsal buccal morphology of a Microhyla rubra tadpole (Stage 35). Scale bar, 1 mm.

Measurements (mm), bh = 5.25; bw = 5.93; ed =

1.26; ht = 5.50; If = 2.54; nn = 1.12; np = 2.66; pp = 5.54;

rn = 1.20; ss = 7.58; su = 7.66; svl = 7.99; tl = 27.04; uf

= 0.85; vt = 19.05; tmh = 2.31, and tmw = 1.68. Mea-

surements of tadpoles in Stages 26, 31, 33, and 35 are

presented in Table 1.

Ecological notes. We observed a group of late-stage

embryos (almost ready to hatch) on the surface of an

open pool of water. The pool was man-made (probably

excavated clay for brick-making forming the depression

which then filled with water), isolated from other water

bodies, and exposed to direct sunlight. The pool shore

was lined with small shrubs and visible submerged ter-

restrial shrubs and vegetation, suggestive of recent in-

undation (Fig. 1). The pool apparently had been filled

with rainwater, and was likely ephemeral. The maximum

depth of the pool was about 50 cm (most areas shallower)

with an area of approximately 100 m^. Water quality of

the pool (9:50 am): temperature = 26.3 °C; dissolved

oxygen = 0.92 mg/1; pH = 6.68; conductivity = 87.8 pS;

salinity = 0; (N 03 )N = 0.524 mg/1; (NH/)N = 0.46 mg/1;

free NH^ = 0.56 mg/1; fiuoride = 0.8 mg/1; total hardness

= 275 mg/1; = 0 mg/1. A total of 410 early stage,

whitish-gray embryos were observed and several were

collected for study.

The larvae of several anuran species were observed

in syntopy with the M. rubra tadpoles: Polypedates mac-

ulatus, Microhyla ornata, Fejervarya limnocharis, a bu-

fonid tadpole of an unidentified species, and Sphaerothe-

ca rolandae.

Discussion

Tadpoles of Microhyla rubra lack keratinized mouth

parts and have a dorsoterminal mouth. Dorsoterminal

mouths are not observed among New World microhy-

lid tadpoles, but within old world microhylid tadpoles,

both terminal and dorsoterminal mouthparts are observed

(Donnelly et al. 1990).

Donnelly et al. (1990) highlighted several microhy-

lids species that lack flaps of the upper lip (M. rubra lacks

flaps on the upper lip) and other species that lack flaps are

Glyphoglossus molossus, Kalaula borealis, K. rugifera,

K. verrucosa, Metaphrynella pollicaris, Microhyla acha-

tina. Mi. anectens. Mi. okinavensis. Mi. heymonsi. Mi.

pulchra, and Mi. zeylanica. Microhyla zelanica is a Sri

Lankan endemic whose tadpole was described by Kir-

tisinghe (1957); though he did not describe the oral flaps

explicitly, his figure shows flaps to be absent on the upper

lip. Kirtisinghe (1957) described tadpoles of M. rubra,

which lack flaps on the upper lip.

Microhyla rubra have six papillae (scallops) on the

lower lip but number varies with developmental stage.

amphibian-reptile-conservation.org

028

December 2011 I Volume 5 I Number 2 I e30

Bowatte and Meegaskumbura

Figure 8. Variation in oral flaps of Microhyla rubra tadpoles at various stages of development (A) Gosner stage 25 - early; (B)

Gosner stage 25 - late; (C) Gosner stage - 30; (D) Gosner stage - 35. Scale bar, 1 mm.

However, in Kirtisinghe’s (1957) diagram of M. rubra,

the scallops are not discernible (not mentioned as papil-

lae or scallops by Donnelly et al. 1990), but there ap-

pears to be more than two, and Kirtisinghe apparently

illustrated a late stage (Stage 35 or later) tadpole. Kirti-

sighe’s (1957) diagram of the lower lip of M. zeylanica

shows five well-distinguished conical papillae. Lower lip

papillae, surprisingly, are reported in few other species of

microhylids (Donnelly et al. 1990).

The whip-like tail-end fiagellum has been reported

from nine species of microhylids (Donnelly et al. 1990).

Parker (1934) and Kirtisinghe (1957) mention the fiagel-

lum in M. rubra. Parker (1934) correctly asserts that the

fiagellum enables these tadpoles to maintain their posi-

tion in water. In aquaria we observed the tail being waved

occasionally but the fiagellum being waved almost con-

tinuously. These tadpoles have the ability to move the

very tail tip, helping maintain their position in the water,

probably helping the tadpoles to conserve energy and

reducing surface disturbance that may be attractive to

predators. Further, buoyancy is perhaps assisted by the

air-filled dorsolateral cavities (or developing lungs) in

the body (in R2).

A nasolacrimal duct is apparent in Stage 35 tadpoles.

Lehr et al. (2007) argue that it is present in all tadpoles,

but only apparent in near metamorphs. Enough informa-

tion has not been gathered to support or refute that this

duct is present in all tadpoles, but it was only apparent

in M. rubra tadpoles at an advanced stage. Lehr et al.

(2007) recommend that a better description for this char-

acter would be to observe whether or not the nasolacri-

mal duct is pigmented. In M. rubra, it is apparent only

because it is relatively unpigmented, compared to the

background, but in some species it may be apparent be-

cause it is more pigmented, compared to the background.

We therefore suggest that when this character is assessed,

the background pigmentation (relative to the pigmenta-

tion on the duct) should be considered.

External nares are open only in late stage microhy-

lid tadpoles (McDiarmid and Altig 1999). Kirtisinghe

(1957) highlights this for M. rubra and we confirm. We

observed that external nares open very late, after front

limbs emerge at Gosner stage 41. Nares opened forming

a rim by the nasal opening in Gosner stage 42.

Kirtishinge (1957) states that toes are fully webbed

in tadpoles. We observed that toes were mostly webbed

in tadpoles (having toes), but saw that webbing rapidly

diminishes by Gosner stage 42. Webbing is vestigial,

conforming to the extent seen in adults, by the one-month

old froglet stage (when the study ended).

The ventral tail fin of M. rubra is deeper than the

dorsal tail fin. Nelson (1972) mentions that Microhyla

amphibian-reptile-conservation.org

029

December 2011 I Volume 5 I Number 2 I e30

Morphology and ecology of tadpoles, Microhyla rubra

Figure 9. Newly emerged froglet of Microhyla rubra (SVL:

8.31mm.

have deeper ventral fins, and highlights M. pulchra and

M. rubra as having much deeper fins. We confirm this

assertion.

The notch apparent on the upper lip, in late stage

(Gosner 35), is not depicted in Kirtisinghe (1957).

The spiracle in M. rubra opens mid-ventrally, and the

opening of the spiracl M. ornata e is crescent-shaped. This

shape is most easily observable in live tadpoles (Fig. 3C).

There is substantial variation in oral flaps at vari-

ous developmental stages (Fig. 8). Most of this variation

is portrayed in the amount and prominence of scallops

on the lower flap (or labium). Variation within Gosner

stages is apparent, especially for early Gosner stages.

For instance, at Gosner stage 25, early-stage larvae have

only two relatively large scallops on each flap, but by

late-stage, size of the individual scallops decreases and

number increases up to six. By Gosner stage 30, number

of scallops remains at six, however, by stage 35, promi-

nence of these are reduced, and in some specimens, de-

pending on the mouth position upon preservation, it can

be difficult to distinguish these scallops. Hence, when

tadpoles are described, it is important to note the devel-

opment of a character periodically over several develop-

mental stages, rather than highlighting characters at only

a single stage (often Gosner stage 35 is used), especially

from only a single individual.

Rao (1918) described M. rubra as being nontrans-

parent, but experience in the field with M. rubra tadpoles

has shown they are almost as transparent as M. ornata

tadpoles. Rao (1918) comments that Ferguson (1904) had

confused the larvae of M. ornata and M. rubra. Howev-

er, without knowing the stage at which the comparisons

were made (there was no general agreement on staging

tadpoles at the time), it is difficult to endorse Rao’s asser-

tion. However, we disagree with Rao’s statement that M.

rubra tadpoles are “not transparent.” Kirtisinghe’s (1957)

description of the Sri Lankan M. rubra refers to them as

“mostly transparent.” However, preservation reduces the

transparency of late-stage tadpoles in both species.

We raised M. rubra for a month beyond metamor-

phosis. This enabled us to determine unequivocally that

the tadpoles raised were verifiably M. rubra (Fig. 9).

Although we sampled for aquatic tadpoles in all hab-

itat types (e.g., man-made irrigation tanks, wells, streams,

rivulets, and paddy fields) we only found M. rubra tad-

poles in ephemeral pools. Several issues could be impor-

tant for their absence: flowing water, water chemistry, the

ephemeral nature of the water body, and predators. The

more permanent water bodies are occupied by predatory

fish such as Channa (Snakehead), Mystus (Catfish), and

smaller cyprinid fishes that we have observed feeding

on the various life history stages of most amphibians. In

these ephemeral habitats, such large aquatic predators are

absent (Skelly 1996; Eterovick and Barata 2006).

Flowing water makes it impossible to have surface-

floating eggs for any length of time. However, the prob-

lem with non-fiowing water is paucity of oxygen, espe-

cially when biomass within the water body is high. One

way of overcoming this is to have surface eggs, which

not only provides for better access to oxygen, but to

higher temperatures, which together facilitate rapid de-

velopment. Rapid development is important when living

in ephemeral pools, to escape desiccation before devel-

opment is complete (Skelly 1996). The temperatures in

the shallow pool (where we found these eggs) were high

(26.3 °C) and oxygen levels low (0.92 mg/1; measured at

9:50 am).

Tadpoles that we raised in the laboratory took 77

days to metamorphose. Days to metamorphose in the

wild might be lower as the temperature in its habitat is

higher (day time lab temperature = 22-24 °C; day time

habitat temperature 26-30 °C), probably accelerating de-

velopment.

M. rubra tadpoles live in water close to the surface

and feed on plankton and suspended food particles.

Many aquatic habitats in the dry zone of Sri Lanka

are polluted to some degree, and ephemeral pools pro-

vide a refuge for amphibians to breed. Activity of the

numerous tadpoles together with the decaying biomass

conceivably could drive up the unbound ammonia and

nitrate concentrations, while reducing the dissolved

oxygen concentration. A combination of indiscriminate

biocide use, overuse of fertilizer, habitat alteration, and

urbanization has changed the freshwater habitats of Sri

Lanka dramatically (Steele et al. 1997). Habitat of early-

phase paddy fields could conceivably provide an excel-

lent environment for M. rubra, although we did not find

them there, conceivably due to the overuse of fertilizer

and biocides. Sri Lanka- Western Ghats is one of the most

populous of the 34 global biodiversity hotspots and this

has created a significant impediment to preserving habi-

tats and moderating rapid changes in inimical land use

patterns.

amphibian-reptile-conservation.org

030

December 2011 I Volume 5 I Number 2 I e30

Bowatte and Meegaskumbura

Water chemistry of the ephemeral pools indicates

that they are not highly polluted. Although free ammo-

nia is fairly high within the pool, bound ammonia (NH^"^)

N, conductivity, salinity, and sulphate-ion concentrations

were low. Further studies are needed to assess the toler-

ance levels of tadpoles and the role of ephemeral pools in

providing a refuge for tadpoles of various species.

Although human activities inadvertently create a few

ephemeral pools for frogs, they may be drained, filled,

and levelled in a surprisingly short period of time. There

is a small chance for breeding populations of frogs to es-

tablish themselves and survive in these types of habitats.

Special consideration (different from those practiced in

preserving and managing the forest habitats of Sri Lanka)

is needed in managing amphibians of the dry zone of Sri

Lanka.

Acknowledgments. — We thank Hendrik Mueller, Ro-

han Pethiyagoda, and Erik Wild for reviewing the manu-

script and providing comments that helped improve the

paper. The following individuals and institutions are

graciously acknowledged: Nimal Gunatilleke and Sav-

itri Gunatilleke for being supportive in numerous ways,

including facilitating transportation; Krishan Ariyasiri

and Udeni Menike for caring for tadpoles; Don Church

and Global Wildlife Conservation for use of equipment

to record water chemistry parameters; James Lewis and

Amphibian Specialist Group for facilitating this work;

and the Department of Wildlife Conservation (DWG) Sri

Lanka for permission to work on tadpoles.

Literature cited

Donnelly MA, de Sa RO, Guyer C. 1990. Description of the

tadpoles of Gastrophryne pictiventris and Nelsonophryne

aterrima (Anura: Microhylidae), with a review of mor-

phological variation in free-swimming microhylid larvae.

American Museum Novitates 2976:1-19.

Eterovick pc, Barata IM. 2006. Distribution of tadpoles with-

in and among Brazilian streams: The influence of predators,

habitat size and heterogeneity. Herpetologica 62(4):365-

377.

Ferguson HS. 1904. A list of Travancore batrachians. Journal

of Bombay Natural History Society 15:505-508.

Gosner KL. 1960. A simplified table for staging anuran em-

bryos and larvae with notes on identification. Herpetologica

16(3):183-190.

GrillitschB, GrillitschH, DuboisA, SplechtnaH. 1 993 . The

tadpoles of the brown frogs Rana {graced) graeca and Rana

igraeca) italica (Amphibia, Anura). Alytes 11(4):117-139.

lUCN. 2004. lUCN Red List Categories and Criteria: Version

3.1. lUCN, Gland, Switzerland and Cambridge, UK. [On-

line]. Available: http://www.iucn.org [Accessed 25 October

2011].

Khan MS. 2000. Buccopharyngeal morphology and feeding

ecology of Microhyla ornata tadpoles. Asiatic Herpetologi-

cal Research 9:130-138.

Kirtisinghe P. 1957. The Amphibia of Ceylon. Published by the

author, Colombo, Sri Lanka. 112 p.

Kirtisinghe P. 1958. Some hitherto undescribed anuran tad-

poles. Ceylon Journal of Science 1:171-176.

Lehr E, Truer L, Venegas PJ, Arbelaez E. 2007. Descriptions

of the tadpoles of two Neotropical microhylid frogs, Mela-

nophryne carpish and Nelsonophryne aequatorialis (Anura:

Microhylidae). Journal of Herpetology 41(4):581-589.

Manamendra-ArachchiK, PethiyagodaR. 2006. Amphibians

of Sri Lanka. Wildlife Heritage Trust, Colombo, Sri Lanka.

440 p.

McDiarmid RW, Altig R. 1999. Tadpoles: The Biology of An-

uran Larvae. The University of Chicago Press, Chicago, II-

linios, USA. 444 p.

Nelson CE. 1972. Systematic studies of the North American

microhylid genus Gastrophryne. Journal of Herpetology

6(2):111-137.

Parker HW. 1928. The brevicipitid frogs of the genus Micro-

hyla. Annals and Magazine of Natural History 2:473-499.

Parker HW. 1934. A Monograph of the Frogs of the Family

Microhylidae. Trustees of the British Museum, British Mu-

seum of Natural History, London, UK. 208 p.

Rao CRN. 1918. Notes on tadpoles of Indian Engystomatidae.

Records of Indian Museum 15:41-45.

Skelly DK. 1996. Pond drying, predators, and the distribution

of tadpoles. Copeia 1996(3):599-605.

Steele P, Konradsen F, Imbulana KAUS. 1997. Irrigation,

health and the environment: A literature review with ex-

amples from Sri Lanka, Colombo, Sri Lanka. International

Irrigation Management Institute (IIMI). Discussion paper

number no. 42. 5:1-25.

Wassersug RJ. 1976. Oral morphology of anuran larvae: Ter-

minology and general description. Publications of Museum

of Natural History, University of Kansas 48:1-23.

Wassersug RJ. 1980. Internal oral features of larvae from eight

anuran families: Functional, systematic, evolutionary and

ecological considerations. Publications of Museum of Natu-

ral History, University of Kansas 68:1-146.

Manuscript received: 28 September 2011

Accepted:23 October 2011

Published: 29 December 2011

amphibian-reptile-conservation.org

031

December 2011 I Volume 5 I Number 2 I e30

Morphology and ecology of tadpoles, Microhyla rubra

GAYAN BO WATTE graduated from the University of Perad-

eniya in 2008. He studied the amphibian diversity changing

with elevation gradient along the Maha-Oya, Hantana forest

for his final year research project and as part of his degree re-

quirements. He is currently a graduate student at the Postgradu-

ate Institute of Science, University of Peradeniya and works

on nitrogen-based stressors affecting amphibians. Gayan also

works on systematics and morphophological development of

tadpoles.

MADHAVA MEEGASKUMBURA is currently a Senior Lec-

turer at the Department of Zoology, Eaculty of Science, Uni-

versity of Peradeniya, Sri Lanka. He is an evolutionary biolo-

gist and ecologist by training and received his B.Sc. in Zoology

from the University of Peradeniya and a Ph.D. from Boston

University (2007). Upon receiving his doctorate degree he was

a Ziff Environmental Postdoctoral Eellow for two years at Har-

vard University (Harvard University Center for the Environ-

ment and Museum of Comparative Zoology). Over the past

decade he has done research on systematics and phylogenet-

ics, evolution, and ecology of Sri Lanka’s frogs, mammals, and

fish. He is the Co-Chairman of the Amphibian Specialist Group

Sri Lanka (ASGSL/IUCN/SSC) and a member of the Amphib-

ian Redlisting Authority (ARLA/IUCN/SSC). Madhava has

published about 20 peer-reviewed papers, several book chap-

ters, and popular articles. Madhava has described about 20 new

species of Sri Lankan animals (frogs, fish, and a mammal) and

a new frog genus (Taruga).

amphibian-reptile-conservation.org

032

December 2011 I Volume 5 I Number 2 I e30

Figure 1. Oligodon arnensis, a non-endemic colubrid snake species found in the lowlands throughout the island, except the dry

southeastern parts. Photo by Indraneil Das.

February 2012 | Volume 5 | Number 2 | e37

amphibian-reptile-conservation.org

033

Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the

original author and source are credited.

Amphibian & Reptiie Conservation 5(2):33-51.

Conservation of biodiversity in a hotspot:

Sri Lanka’s amphibians and reptiies

WALTER R. ERDELEN

115, route du Tertre, 91530 Sermaise, FRANCE

Abstract . — Sri Lanka is a continental tropical island that is considered a hotspot for amphibian and

reptile diversity. During the last decade herpetological research has substantially improved our

knowledge of species and their taxonomic status. However, additional work is needed on ecology

and population viability within the framework of human impacts on natural ecosystems. These hu-

man induced activities have led to severe fragmentation of formerly continuous forest in the wet

zone and central hills of Sri Lanka, where most endemic and threatened species occur. Here I dis-

cuss current development in biodiversity issues regarding the Convention on Biological Diversity

and their effects on the future of herpetofaunal conservation in Sri Lanka. To better understand Sri

Lanka’s conservation challenges and threats I discuss the following topics: Sri Lanka’s biogeogra-

phy; its extant ecosystems and landscapes along with the changes resulting from patterns of hu-

man settlement; human population growth and its concomitant impact on natural ecosystems; and

a brief history of herpetological studies in Sri Lanka. Further, I discuss major conservation issues

related to the ecoregional and hotspot approach to biodiversity conservation, the lUCN species

lists, and the institutional framework in biodiversity conservation. Finally, I propose an integrated

action plan for the conservation of Sri Lanka’s herpetofauna that includes cooperation between

relevant institutions, future scientific studies, education, capacity development, in situ and ex situ

conservation, and encouragement of increased collaborative effort in biodiversity conservation

with the Western Ghats of southern India.

Key words. Sri Lanka, biogeography, history of herpetological research, biodiversity conservation, biodiversity

hotspot, amphibians, reptiles, action plan

Citation: Erdelen WR. 2012. Conservation of biodiversity in a hotspot: Sri Lanka’s amphibians and reptiles. Amphibian & Reptiie Conservation 5(2):33-

51 (e37).

Introduction

The World Summit on Sustainable Development, held in

Johannesburg in 2002, and the United Nations General

Assembly endorsed a “2010 Target” based on a decision

of the 6* Conference of the Parties to the Convention on

Biological Diversity. The target was to achieve, by 2010,

a significant reduction of the current rate of biodiversity

loss at global, regional, and national levels as a contribu-

tion to poverty alleviation and to the benefit of all life

on Earth (SCBD 2010). The 2010 target and its 21 sub-

targets have not been met globally despite partial local

achievements (SCBD 2010). To scale up efforts to deal

with continued biodiversity loss and other biodiversity

issues the United Nations proclaimed 2010 the “Interna-

tional Year of Biodiversity.” The main objectives of the

Year were to (source: Secretariat of the Convention on

Biological Diversity):

• Enhance public awareness of the importance of con-

serving biodiversity and underlying threats to

biodiversity.

• Raise awareness of accomplishments to save biodi-

versity by communities and governments.

• Promote innovative solutions to reduce threats to

biodiversity.

• Encourage individuals, organizations, and govern-

ments to take immediate steps to halt biodiversity

loss.

• Initiate dialog between stake holders for steps taken

in the post-2010 period.

In October 2010 the 10* meeting of the Conference of the

Parties to the Convention on Biological Diversity (COP

10) took place in Nagoya, Japan. Efforts in Nagoya were

Correspondence. Email: walter.erdelen® gmail.com

February 2012 | Volume 5 | Number 2 | e37

amphibian-reptile-conservation.org

034

Erdelen

underpinned by earlier reports on biodiversity such as the

biodiversity synthesis report of the Millennium Ecosys-

tem Assessment (MEA 2005) and Global Biodiversity

Outlook 3 (SCBD 2010). The COP 10 meeting was a

breakthrough in the conservation of biological diversity.

Meeting participants adopted an outstanding measures

package including: (1) a strategic plan for biodiversity

and the Aichi biodiversity targets; (2) the Nagoya pro-

tocol on access to genetic resources and fair and equi-

table sharing of benefits arising from their utilization; (3)

a strategy for resource mobilization; (4) a continuation

of the process of establishing an intergovernmental plat-

form on biodiversity and ecosystem services; and (5) the

recommendation to the United Nations General Assem-

bly to declare 2011-2020 the UN Decade on Biodiversity.

One key outcome of the COP 10 meeting was the

recommendation to globally update the national biodi-

versity strategies and action plans (NBSAPs). Within the

process of updating, amphibians and reptiles could get

more attendance within the overall framework of pre-

serving Sri Lanka’s unique biodiversity. The relevance

of an adequate consideration of Sri Lanka’s herpetofauna

for NBSAP is that Sri Lanka is recognized as a global

amphibian hotspot (Meegaskumbura et al. 2002; Pethi-

yagoda and Manamendra-Arachchi 1998) as well as a

mega-hotspot of reptile diversity (Somaweera and So-

maweera 2009).

Moreover, especially since the release of the 4*

Assessment Report of the IPCC (2007; see: www.ipcc.

ch) and the so-called “Stern Review” (Stern 2006), the

global political leadership and the UN have increasingly

focused on discussions of global climate change and its

effects on human well-being and the future of Earth’s

biological diversity. Collectively these most recent de-

velopments seem to set the stage for new discussions

about conserving Sri Lanka’s biodiversity and mitigat-

ing the impacts of — and adapting to — global climate

change. The herpetofauna of Sri Lanka, being an essen-

tial component and an indicator of the overall health of

Sri Lanka’s ecosystems, plays a crucial role in contrib-

uting both to the sustenance of the country’s wealth in

life forms and ecosystem services provided to the local

human population.

This paper is future-oriented and action-oriented

with regard to the long term preservation of Sri Lanka’s

herpetofauna. Here I provide a holistic picture of what

is needed to strengthen conservation efforts at all levels,

including research, education, partnership, and policy.

These conservation efforts should be accomplished first

and foremost at the national level but also integrated

into subregional (e.g., jointly for the Western Ghats of

India and Sri Lanka biodiversity hotspots), regional, and

global efforts toward amphibian and reptile conserva-

tion. These conservation efforts should be recognized

in context to human impact on natural ecosystems and

global climate change. Moreover, they should be part of

Sri Lanka’s overall effort towards biodiversity conserva-

tion and sustainable use of its ecosystem services (for an

overview see TEEB 2010). More specifically, this paper

outlines: (1) aspects of the biogeography of Sri Lanka;

(2) the history of herpetological research and our current

knowledge base; (3) conservation issues; and (4) a pro-

posal intended to contribute to further discussions and

elicit appropriate measures for future sustainable conser-

vation of Sri Lanka’s herpetofauna.

The tropical continental island of Sri

Lanka — A note on biogeography

Historical remarks

Based on detailed studies of the fiora and fauna of India

over thirty-five years ago, attempts were made to sub-

divide the Indo-Ceylonese region into biogeographical

subregions and other units (e.g., Mani 1974). The first

zoogeographical studies, carried out in the 19* century,

were based on distributional patterns of terrestrial mol-

lusks (Blanford 1870), reptiles (Gunther 1858, 1864),

and birds (Jerdon 1862-1864). The definition of fioris-

tic regions began in the middle of the 19* century (e.g..

Hooker and Thomson 1855; Clarke 1898) and the begin-

ning of the 20* century (e.g., Prain 1903; Hooker 1906).

Collectively, these studies revealed a strong similar-

ity between Sri Lanka and neighboring India, especially

with regard to the more humid regions of the Western

Ghats and southwestern Sri Lanka. Repeatedly, south In-

dia and Sri Lanka were seen as a single biogeographical

subunit comprising two major pairs of similarities, i.e.,

the Malabar Tract, southwestern and hill regions of Sri

Lanka, southeastern India, and drier parts of Sri Lanka

(e.g., Bhimachar 1945; Phillips 1942; Wait 1914). These

patterns of similarity encompass the majority of plant

and animal species, particularly the herpetofauna dis-

cussed here (for an overview of the biogeography of the

reptiles of south Asia, see Das 1996a).

Geological past

The geological history of Sri Lanka is subdivided into

the following phases (after Dietz and Holden 1970; Keast

1973; McKenna 1975; Pielou 1979; Raven and Axelrod

1974):

• Pre-drift phase where Sri Lanka and India were part

of Gondwana (> 100 MYBP).

• Drift phase ending with the collision of the In-

dian plate and the Asiatic continent (66 and 45

MYBP).

• Miocene epoch (ca. 25 MYBP), Sri Lanka’s sepa-

ration from India, following a series of complex

February 2012 | Volume 5 | Number 2 | e37

amphibian-reptile-conservation.org

035

Sri Lanka’s amphibians and reptiles

tectonic movements, which began in the Jurassic

(see Cooray 1984; Katz 1978; Swan 1983).

• Quaternary epoch (two MYBP to present), eustatic

sea level changes, climate cycles, and repeated

formation of land bridges between India and Sri

Lanka, in the Palk Strait region.

Similarities observed between flora and fauna of Sri

Lanka and India are linked to having been part of the

Indian plate and an isolated unit in the Tethys Sea, after

its separation from the Gondwanan landmass and before

it collided with Asia. Additionally, the biogeographical

evolution of India and Sri Lanka was certainly shaped by

the global K-T event, the Deccan volcanism (Cretaceous

to Eocene; Wadia 1976), the orogenic processes leading

to formation of the Himalayas, the development of the

monsoon pattern, and floristic and faunistic exchanges

between the Indian plate and Asia (early Tertiary 45-25

MYBP), particularly with southeast Asia (see Klaus et al.

2010). This phase was followed by Quaternary climate

fluctuations and eustatic changes in sea level leading to

repeated formation of land bridges between India and

Sri Lanka (Palk Strait region; for pollen data see Prema-

thilake and Risberg 2003). During Quaternary sea level

maxima, when Sri Lanka was isolated from India, bio-

geographical patterns most likely changed independently

from India. The Quaternary is often seen as the decisive

period for shaping the present plant and animal distribu-

tion patterns in Sri Lanka (e.g., Erdelen 1993a; Erdelen

and Preu 1990a). “Time lags” between eustatic sea lev-

el changes, climate change, and the “reaction” of plant

and animal species may explain some of the similarities

among rain forest species in southern India and Sri Lanka

(Erdelen and Preu 1990a).

Many unanswered questions exist regarding the bio-

geographical evolution of Sri Eanka’s flora and fauna

(for more recent analyses see Biswas 2008; Biswas and

Pawar 2006). Most speciation events among amphib-

ians and reptiles pre-date the Quaternary period. This

notion is supported by several recent papers on genetic

divergence within rhacophorid frogs. A study on rostral

horn evolution of the endemic genus Ceratophora sug-

gests a Miocene origin of the genus and several specia-

tion events dating approximately between 12.6 and 2.4

MYBP (Schulte II et al. 2002). A similar situation was

reported for the remarkable radiation of Sri Lanka’s

freshwater crabs (50 endemics from a total of 51 species

for the island; Beenaerts et al. 2010). The uropeltid snake

species of southern India and Sri Lanka may have been

separated for a period longer than 10-15 MYBP (e.g.,

Cadle et al. 1990). In fact, many of the speciation events

thought to have been associated with different phases of

the Pleistocene are much older and likely the result of

speciation events in the Tertiary (e.g., see Maxson 1984,

Roberts and Maxson 1985a, 1985b, for Australian frogs).

Speciation rates may have varied within groups such

as birds in Sri Lanka and India (Erdelen 1993a). Migra-

amphibian-reptile-conservation.org

tion patterns into and out of the Indian-Sri Lankan region

likely differed substantially among and within taxa (for

Cincidelid beetles, see Pearson and Ghorpade 1989), and

exchanges of floral and faunal elements need not have

been symmetric but may show a marked asymmetry if

India and neighboring regions are compared. The results

of these highly variable processes are rather complex ex-

tant patterns of geographic distribution. Eurther studies

are essential for a more complete understanding of the

major evolutionary processes that formed Sri Lanka’s

flora and fauna. The basis of such studies would be the

understanding of undisturbed, “pristine” geographic dis-

tribution patterns allowing for the reconstruction of his-

torical processes producing Sri Lanka’s biodiversity.

Extant ecosystems and landscapes

Sri Lanka’s rich biodiversity is reflected in its diverse

extant ecosystems and landscapes. Ecosystems may be

classified into the following (for more details and refer-

ences, see Dela 2009; Gunatilleke et al. 2008; Ministry of

Eorestry and Environment 1999):

• Forest and grassland

• Inland wetland

• Coastal and marine

• Agricultural

• Urban

The most important ecosystems for amphibians and rep-

tiles are certainly the first two categories, especially if

minimally disturbed by humans, although coastal and

marine ecosystems are important to reptile taxa like ma-

rine turtles and crocodiles. Agricultural and urban sys-

tems may provide habitats for species with broad habitat

requirements, especially those that live commensally

with humans.

Often underestimated in their role of maintaining

viable populations are secondary forests or, more gener-

ally, “novel ecosystems.” These are described as heavily

influenced by humans but not under human management,

or “lands without agricultural or urban use embedded

in agricultural and urban regions” (Marris 2009). More

than 90% of amphibian species in Sri Lanka occur in

secondary forests, highlighting the importance of novel

ecosystems (R. Pethyiagoda, pers. comm.). Long-term

conservation efforts should consider the landscape mo-

saic of Sri Lanka, which comprises ecosystems that vary

in geographic extent and human perturbation. System

interlinkages and scale may be essential parameters for

understanding and managing such diverse environments

(Erdelen 1993b).

Vegetation maps for Sri Lanka date to the 1930s.

Based on the three climatic zones of the island, namely

the wet, intermediate, and dry zones, the National Atlas

of Sri Lanka distinguished 11 different types of plant

communities (Somasekaram 1988). For analyses of fau-

February 2012 | Volume 5 | Number 2 | e37

036

Erdelen

nal distribution patterns in Sri Lanka a simplified subdi-

vision into seven zones with six different types of natural

vegetation has been frequently used (e.g., Crusz 1984,

1986; Crusz and Nugaliyadde 1978; Erdelen 1984, 1989,

1993a).

Based on distribution data for angiosperm plants,

recent studies have shown that within these major veg-

etation units 15 fioristic regions may be distinguished,

located largely within the wet zone and the mountain re-

gion of Sri Lanka (Ashton and Gunatilleke 1987; Guna-

tilleke and Gunatilleke 1990). Even within these fioristic

regions, forest communities show a patchy distribution,

sometimes with rather different species compositions

(Gunatilleke and Gunatilleke 1983). Individual hills may

have unique forest communities (Abeywickrama 1956),

for example Hinidumkande in the southwestern part of

the wet zone. The rainforests of this mountain show a

striking concentration of endemic tree species (Guna-

tilleke and Gunatilleke 1984). Another well-known ex-

ample is Ritigala, a 766 m high mountain in the northern

part of Sri Lanka’s dry zone. Although located in the dry

zone this mountain contains endemic plant species char-

acteristic of the wet zone and species which otherwise

occur only in the mountain region and not elsewhere in

the dry zone. Some plant species are endemic to Ritigala

(for details see Jayasuriya and Pemadasa 1983; Jayas-

uriya 1984).

Although numerous attempts have been made to ex-

plain these highly localized concentrations of endemic

species (see Willis 1916, for one of the earlier discus-

sions), we still do not know whether, and to what ex-

tent, these are possibly a result of Quaternary dynamics

of vegetation patterns (related to glacial and interglacial

cycles and associated climate regimes). Moreover, it is

not clear whether, and if so to what extent, such small-

scale mosaics in vegetation patterns are refiected in en-

demic animal taxa, and thus may need more attention as

part of the overall efforts of biodiversity conservation in

Sri Lanka (see Raheem et al. 2009).

When we try to reconstruct the evolution of Sri Lan-

ka’s biota and its relationship to Indian fiora and fauna,

“biogeographical reconstruction” is increasingly ham-

pered by anthropogenic alterations of habitats. Relatively

undisturbed ecosystems and associated distribution pat-

terns within a fioral or faunal setup should be the basis

for reconstructing historical events, which shaped the

extant composition of Sri Lanka’s fiora and fauna. Only

if the spatio-temporal dynamics of anthropogenic effects

on natural ecosystems are well-known and documented

will such a reconstruction process be facilitated and the

“true” patterns and underlying historical processes in-

volved be discovered.

Modem humans settled in Sri Lanka between 75,000

and 125,000 YBP or earlier (Deraniyagala 1993). Esti-

mates of human densities during different periods of

human history in Sri Lanka would provide indirect evi-

dence of potential impacts on natural vegetation and as-

sociated fauna. During the pre-historic phase, between

75,000 YBP and 10,000 YBP, when humans were es-

sentially subsistence hunters and food gatherers, the wet

zone and hills of Sri Lanka were already settled, although

in low densities. Deraniyagala (1993) provides an esti-

mate for the wet zone during this phase of up to 10,000

YBP of some 0.1 individuals/km^. The transition period

(pre-historic to proto-historic and early historic phases),

saw high human densities in the dry zone increasing dur-

ing the Singhalese high culture (beginning ca. 200 BC),

a time associated with the advent of Buddhism in Sri

Lanka. During the Anuradhapura Period (250 BC-1017;

first urbanization phase) and the Polonnaruwa Period

(1017-1235) extensive systems of irrigation tanks were

established in the dry zone for rice cultivation (see Abey-

wickrama 1993).

During the Late Historic Phase, from the 14* centu-

ry onwards, the political, economic, and cultural centers

shifted from the north-central, eastern and southeastern

parts of the island towards the lowlands of the wet zone,

the central highlands, and into the extreme northern parts

of Sri Lanka (Erdelen 1993a). This restmcturing process

was associated with the downfall of high cultures in the

dry zone and the beginning of the colonial periods (Por-

tuguese, Dutch, and British). During the British Period

(1796-1948) in particular, massive impacts on the natural

forests of southwestern Sri Lanka and the central hills

were recorded. The introduction of plantation industry

(cinchona, coffee, tea, and mbber) and infrastmctural

measures caused changes for these regions. Eollowing

Sri Lanka’s independence (1948), there was a period of

intensified man-made alterations to the natural ecosys-

tems of Sri Lanka, with the objective of supporting both

a rapidly increasing population and an accelerated eco-

nomic growth (Erdelen 1988b, 1993; Erdelen and Preu

1990b; Erdelen et al. 1993; Ministry of Eorestry and En-

vironment 1999).

The population of Sri Lanka has tripled in size in

some 60 years, from 7.2 million inhabitants in 1948 to

over 21 mill ion in 2011. Population density, formerly be-

ing highest in the dry zone of Sri Lanka, has now reached

over 500 individuals/km^ in the wet zone (Dela 2009;

see Cincotta et al. 2000, with regard to global biodiver-

sity hotspots). These historical processes have led to a

considerable change in the distribution of natural veg-

etation in Sri Lanka (see Erdelen 1996). More extensive

areas under natural forest cover are essentially found in

the dry zone. The forests of the wet zone and the central

hill range have become highly fragmented. No continu-

ous primary forest cover remains from sea level to over

2,500 m of the central hill range. Note these statements

refer to “vegetation” and major types of ecosystems but

do not refiect the fine-scale analysis and implications

these changes might have for plant and animal species/

populations and their long-term viability.

February 2012 | Volume 5 | Number 2 | e37

amphibian-reptile-conservation.org

037

Sri Lanka’s amphibians and reptiles

Analysis of the following questions may be useful

in gaining a better understanding of processes at relevant

scales and for subsequent appropriate conservation mea-

sures:

1) Concomitant with anthropogenic impacts on natu-

ral vegetation: have plant communities changed

significantly both in structure, and therefore, in

microhabitat and microclimatic conditions, as

well as in species composition?

2) If so, at what scale has this happened and what does

the extant mosaic of differentially impacted plant

ecosystems look like?

3) How do distribution patterns of amphibians and

reptiles relate to vegetation or plant communi-

ty patterns? If they do, what is the “reference”

equivalent with regard to vegetation type or

“structural” habitat parameters against which dis-

tribution patterns could be calibrated?

4) What are the projections of population or species

viabilities if questions 1-3 are analyzed simulta-

neously?

5) What would be the implications of such analyses

for biodiversity conservation measures, specifi-

cally in regards to amphibians and reptiles?

In conclusion, we need a better understanding of proxi-

mate and ultimate factors (i.e., knowledge of the crucial

ecosystem or habitat parameters) decisive in the long-

term persistence of amphibian and reptile populations.

These factors vary intrinsically with species’ ecologies

and are shaped by human impacts on natural ecosystems

and habitats. These concepts need to be taken into ac-

count for monitoring long-term population trends in Sri

Lanka.

History of herpetological research

in Sri Lanka

Herpetological research has a long history in Sri Lanka

(de Silva 2001) and has been part of the general history

of biodiversity exploration in Sri Lanka (Pethiyagoda

2007). Interest during the British period (1796-1948)

was mainly in horticulture for the introduction of com-

mercially-used crops and for exporting plants from Sri

Lanka. Except for earlier work by French workers and

scientists associated with the British Museum in the 19*

century, the focus on the fauna of Sri Lanka began with

the establishment of the Colombo Museum in 1877. For

the most part, until about the time of independence, it

would be amateurs who led efforts to explore the island’s

herpetofauna (Pethiyagoda 2007).

A detailed analysis of factors shaping herpetological

research in Sri Lanka would be worth undertaking but is

beyond the scope of this paper. The most recent scientific

research efforts have been vital for a more thorough un-

derstanding of the herpetofauna of Sri Lanka, especially

in regard to the number of species on the island as well as

their taxonomic status. It is clear from these studies that

several species have become extinct in recent times and

more work is needed to preserve Sri Lanka’s herpetofau-

nal diversity into the future (see below).

Amphibians

Species lists for amphibians of Sri Lanka have been com-

piled since the 19* century. These were first published

within the framework of regional compilations such as

the works of Gunther (1864) and Boulenger (1890). The

first lists of exclusively Sri Lankan amphibians were

published by Kelaart (1852) and Haly (1886a) followed

by numerous publications on individual amphibian taxa

(for compilations see Dutta and Manamendra-Arachchi

1996; Erdelen 1993a). In the 1950s, de Silva published

a species list for Sri Eanka, including the specimens

housed in the Colombo Museum (de Silva 1955). This

Figure 2. Tadpoles (top) and adult specimen (bottom) of

Nannophrys marmorata, an endemic species restricted to the

Knuckles range; Critically Endangered. Mainly found under

boulders on wet, flat, rocky surfaces (Dutta and Manamendra-

Arachchi 1996; confirmed by own observations). The genus is

endemic to Sri Lanka, comprising four species, one of them

(N. naeyakai) described only in 2007 (Fernando et al. 2007).

Photos by Walter R. Erdelen.

February 2012 | Volume 5 | Number 2 | e37

amphibian-reptile-conservation.org

038

Erdelen

publication was followed by Kirtisinghe’s (1957) mono-

graph The Amphibia of Ceylon. Thereafter, and repeat-

edly, checklists for the amphibians of Sri Lanka were

compiled (Kotagama et al. 1981; de Silva 1994, 1996,

2001). In parallel, taxonomic revisions were undertaken

for the first time (for details see Dutta and Manamendra-

Arachchi 1996 and Erdelen 1993a). Dutta (1985), in his

Ph.D. dissertation, updated information on the amphib-

ians of Sri Lanka and India and in 1996 published the

first modern account of the amphibian fauna of Sri Lanka

(Dutta and Manamendra-Arachchi 1996). Possibly the

first indication that Sri Lanka may be home to many more

amphibian species is indicated in publications from the

mid-90s where new amphibian species were described

(e.g., Fernando et al. 1994; Manamendra-Arachchi and

Gabadage 1996). As Dutta and Manamendra-Arachchi

(1996) wrote in their introduction: “We expect there to

be a dramatic increase in the diversity of amphibians of

Sri Lanka, especially among the Rhacophoridae.” Indeed

in 2002 detailed information on Sri Lanka’s outstanding

amphibian diversity was published in an article in Science

(Meegaskumbura et al. 2002) indicating that rhacophorid

frogs may comprise over 100 species in Sri Lanka. In this

paper it was stated that “Sri Lanka’s amphibian diver-

sity (about 140 species on an island of 65,610 km^) now

approaches or exceeds that of many amphibian diversity

hotspots and is comparable to those of tropical islands

an order of magnitude larger, such as Borneo (746,300

km^; 137 species), Madagascar (587,000 km^; 190 spe-

cies), New Guinea (775,200 km^; 225 species), and the

Philippines (299,800 km^; 96 species).”

Meanwhile, species numbers for amphibians in Sri

Lanka stand at 111, of which some 90% are endemic

(Fig. 2; for regularly updated information see: http://am-

phibiaweb.org). Still more species await description and

the percentage of endemism is expected to rise, as seen

in the 2007 list of threatened fauna and fiora of Sri Lanka

which already mentions 106 amphibian species of which

90 (85%) are endemic (lUCN Sri Lanka and MoENR

2007).

Reptiles

The earliest publications on Sri Lankan reptiles are in-

cluded in those of a more general nature already men-

tioned above. Ferguson (1877) and Haly (1886b, 1891)

compiled information about reptiles in collections of the

Colombo Museum. Most famous have been the publica-

tions of P. E. P. Deraniyagala (for an overview, see de Sil-

va 1977). He published three outstanding volumes on the

turtles and crocodiles, lizards, and snakes of Sri Lanka

(Deraniyagala 1939, 1953, 1955). At that time, the only

comparable publications were Smith’s Fauna of British

India (Smith 1931, 1935, 1943) and Taylor’s work on in-

dividual taxa (Taylor 1947, 1953b) and his overviews of

the Sri Lankan snakes, skinks, and lizards (Taylor 1950a,

1950b, 1953a).

This period was followed by a number of system-

atic/taxonomic and ecological studies of individual taxa

(overviews in Erdelen 1993a; de Silva 2006). De Silva

(1998a, 1998b, 1998c) published checklists and anno-

tated bibliographies of the turtles and crocodiles, lizards,

and snakes of Sri Lanka. Comprehensive publications

are available on snakes (de Silva 1980) and color guides

were more recently published on snakes (de Silva 1990)

and lizards (Somaweera and Somaweera 2009) of Sri

Lanka.

The 2007 Red List of Threatened Fauna and Flora

of Sri Lanka (lUCN Sri Lanka and MoENR 2007) lists a

total of 171 reptile species where 101 (59%) are endemic

(Fig. 3), with more being added (e.g., Gower et al. 2011;

Maduwage et al. 2009).

The herpetofauna of Sri Lanka — A short

summary of the evolution of our knowledge

base

Although our knowledge of Sri Lankan herpetofauna

has considerably improved, new species still await dis-

covery. This applies particularly to amphibians where

traditional morphological approaches have fallen short

of adequately describing species diversity (for compari-

son see Oliver et al. 2009; Stuart et al. 2006; Vieites et

al. 2009). Modern genetic analyses have shown a much

higher species diversity than previously expected (over-

view in Pethiyagoda et al. 2006). In addition, new species

of reptiles have been discovered during the last years of

intensified field work in Sri Lanka. This includes “seem-

ingly” better known agamid genera such as Calotes, Cer-

atophora, Cophotis, and Otocryptis (for an overview, see

references in Bahir and Surasinghe 2005 and Somaweera

and Somaweera 2009; Fig. 4). In addition, new species

of scincid and gekkonid lizards and snakes were recently

Figure 3. Male specimen of Lyriocephalus scutatus, the most

charismatic lizard of Sri Lanka. The genus is monotypic and

endemic to Sri Lanka. Photo by Walter R. Erdelen.

February 2012 | Volume 5 | Number 2 | e37

amphibian-reptile-conservation.org

039

Sri Lanka’s amphibians and reptiles

described (overviews in de Silva 2006; Somaweera and

Somaweera 2009).

As already indicated by Pethiyagoda et al. (2006),

despite recent work on taxonomy and systematics com-

paratively little is known about the biology of Sri Lankan

amphibians. Basic ecological information at both the

population and species levels is unavailable for most,

if not all taxa. Additionally, geographic distribution

patterns and their dynamics are poorly understood or

not known at all. The rarity of amphibian species, their

patchy distribution, and possibly highly fragmented or

small populations have neither been adequately recorded

nor monitored over time, especially in view of human-in-

duced habitat or microhabitat changes. Similarly, we lack

this information for most Sri Lankan reptile species as

well. An exception may be studies on the genus Calotes

including analyses of geographic distribution patterns,

intraspecific variability, and population dynamics (Erdel-

en 1977, 1983, 1984, 1988a; for a more recent study of

C. nigrilabris see Amarasinghe et al. 2011).

Our knowledge of amphibian and reptile diversity in

Sri Lanka has profoundly improved during recent times

(within the last decade). This improvement has been the

result of a “new age of herpetology, characterized both

by increased international cooperation in research and by

the blossoming of herpetology as a research discipline

for many young Sri Lankan zoologists” (de Silva 2006).

This process was infiuenced or catalyzed by major her-

petological events held in Sri Lanka, including the 1996

International Conference on the Biology and Conserva-

tion of the Amphibians and Reptiles of South Asia, held

at the University of Peradeniya (de Silva 1998), and the

4 th World Congress of Herpetology, held at Bentota, Sri

Lanka in 2001 (see Dodd and Bartholomew 2002).

Conservation issues

Generai observations

Sri Lanka has a long tradition of preserving its wildlife.

It was one of the earliest countries to set aside areas for

wildlife protection and take conservation measures for

its plant and animal life. Ideas of preserving nature in Sri

Lanka may date back to the advent of Buddhism, about

2,500 YBP. Sanctuaries were already established in Sri

Lanka in the 12* century, possibly earlier (see Cmsz

1973; DeAlwis 1969; Erdelen 1988b; Ministry of For-

estry and Environment 1999).

Currently, Sri Lanka has over 500 protected areas in-

cluding over 90 key biodiversity areas recently identified

jointly by the Wildlife Heritage Trust and the University

of Peradeniya. Sri Lanka’s protected areas — covering

about 18% of the island’s total land area — are principally

Figure 4. Range restricted endemic forest lizards. Top left: Ceratophora tennentii, male; top right: Cophotis ceylanica, male; bot-

tom left: Calotes liocephalus, juvenile; bottom right: a newly discovered endemic but widespread species of scincid lizard {Eutropis

tammanna', described by Das et al. 2008). Eutropis tammanna photo by Indraneil Das; all others by Walter R. Erdelen.

February 2012 | Volume 5 | Number 2 | e37

amphibian-reptile-conservation.org

040

Erdelen

Figure 5. Two species of reptiles endemic to the Knuckles range, the gekkonid Cyrtodactylus soba (left) and the scincid Nessia

bipes (right). Photos by Indraneil Das.

managed by the Forest Department and the Department

of Wildlife Conservation (for details see Dela 2009).

The most recent significant international achievement

has been the recognition of the Central Highlands of Sri

Lanka, including the Peak Wilderness Protected Area,

the Horton Plains National Park, and the Knuckles Con-

servation Forest (see Fig. 5), as a World Heritage Site.

As stated in the relevant text of the World Heritage

Committee (34 COM8B.9) decision: “the property in-

cludes the largest and least disturbed remaining areas of

the submontane and montane rain forests of Sri Lanka,

which are a global conservation priority on many ac-

counts. They include areas of Sri Lankan montane rain

forests considered as a super-hotspot within the Western

Ghats and Sri Lanka biodiversity hotspot. More than half

of Sri Lanka’s endemic vertebrates, half of the coun-

try’s endemic fiowering plants and more than 34% of its

endemic trees, shrubs, and herbs are restricted to these

diverse montane rain forests and adjoining grassland

areas.” In the same text it is further noted that: “Of the

408 species of vertebrates, 83% of indigenous fresh wa-

ter fishes and 81% of the amphibians in Peak Wilderness

Protected Area are endemic, 91% of the amphibians and

89% of the reptiles in Horton Plains are endemic, and

64% of the amphibians and 51% of the reptiles in the

Knuckles Conservation Forest are endemic.”

As indicated above, conservation efforts in Sri Lan-

ka previously focused largely on charismatic and well-

known species such as the larger mammal and bird spe-

cies and endemic plant and animal species. Amphibians

and reptiles have largely been ignored, a situation similar

to other Asian countries such as Indonesia (Iskandar and

Erdelen 2006). This fact underscores the importance of

specific mention of amphibians and reptiles in the nomi-

nation of this new World Heritage Site, which is of out-

standing importance to the long-term conservation of a

significant segment of Sri Lanka’s herpetofauna and its

fauna and fiora in general.

Sri Lanka’s fourth country report to the Convention

of Biological Diversity lists the following major threats

to Sri Lanka’s biodiversity: (1) habitat loss and frag-

mentation, in particular regarding wet zone ecosystems;

(2) habitat degradation; (3) overexploitation of biologi-

cal resources; (4) loss of traditional crop and livestock

varieties and breeds; (5) pollution; (6) human-wildlife

confiicts; (7) spread of alien invasive species; and (8)

increasing human population density (Dela 2009). With-

out doubt numbers one and two above are the most im-

portant direct threats to the herpetofauna of Sri Lanka,

particularly in regards to endemic species. Pesticide use

and air pollution possibly affect amphibian populations

more drastically than reptiles, due to their complex life

histories (Ariyasiri et al. 2011). The long-term viability

of amphibian populations critically depends on the state

of both the aquatic ecosystems they use during their “bi-

modal” life cycle and the associated terrestrial ecosys-

tems they inhabit (see Becker et al. 2007).

As pointed out by Pethiyagoda et al. (2006), the area

of greatest concern for amphibians is the southwestern

region of Sri Lanka where over 95% of forest cover has

been lost and amphibian species are restricted in their

geographic distribution. The wet zone of Sri Lanka cur-

rently comprises well over 100 forest fragments, and

areas where continuous forest exists from lowlands to

higher elevations are rare. This situation is further ag-

gravated by high human population density in the south-

western region of Sri Lanka with over 500 individuals/

km^ (Dela 2009; see above).

Ecoregions and hotspots of

biodiversity — The case of Sri Lanka

In their paper “Global 200,” Olson and Dinerstein (1998)

identified the 200 biologically most valuable ecoregions.

The terrestrial ecoregions are defined as relatively large

units of land containing a distinct assemblage of natural

communities and species, with boundaries that approxi-

mate the original extent of natural communities prior to

major land-use change (Olson et al. 2001). Biological

distinctiveness was measured in terms of species rich-

ness, endemism, taxonomic uniqueness, unusual eco-

February 2012 | Volume 5 | Number 2 | e37

amphibian-reptile-conservation.org

041

Sri Lanka’s amphibians and reptiles

logical or evolutionary phenomena, and global rarity of

habitat types (for details see Olson and Dinerstein 1998).

This included the moist forests of the Western Ghats and

Sri Lanka — both classified as Critical or Endangered

as their conservation status. A more detailed analysis

was presented in the Indo-Pacific terrestrial ecoregions

conservation assessment (Wikramanayake et al. 2002).

This assessment provided a detailed subdivision of the

Western Ghats and also distinguished three ecoregions

within Sri Lanka: (1) lowland rain forests, (2) montane

rain forests, and (3) evergreen forests of the dry zone.

The first two were considered globally outstanding with

a conservation status of “critical” and given the highest

assessment of need for effective biodiversity conserva-

tion - “class I” (see Fig. 6). The third was classified as

regionally outstanding, vulnerable, and assigned “class

11” as its conservation assessment (for details, see Wikra-

manyake et al. 2002).

In parallel, the assignment of global conservation

priorities was based on the concept of “biodiversity

hotspot,” a term coined by Myers in the late 1980s (My-

ers 1988, 1990). The term originally referred to areas

where “exceptional concentrations of endemic species

are undergoing exceptional loss of habitat” (Myers et al.

2000). Other definitions include parameters like species

richness, degree of endemism, numbers of rare or threat-

ened species, and intensity of threat (see Reid 1998). One

persistent discordant issue is that rare species may not

occur in the most species-rich areas (e.g., Prendergast et

al. 1993; see also Reid 1998; for vascular plant diversity

and hotspots see discussions in Kiiper et al. 2004; Mutke

and Barthlott 2005; Mutke et al. 2011).

Early work described the Western Ghats and Sri

Lanka as a single unit in the list of global biodiversity

hotspots (e.g., in Myers 1990). Based on the following

factors: endemic plant species, endemic vertebrates, the

occurrence of endemic plant and vertebrate species per

100 km^, and the percentage of remaining primary veg-

etation, Myers et al. (2000) identified the “eight hottest

hotspots” and included the Western Ghats and Sri Lanka.

The relationship between the hotspot and ecoregion

approaches is not further discussed here (see e.g.. Ladle

and Whittaker (2011) for discussions of the two ap-

proaches) but a short comment on their interrelationships

is of benefit. Regarding scale, the ecoregional approach

generally is more fine-scale in nature. For instance, the

Western Ghats and Sri Lanka comprise eight different

ecoregions. In general, there is over 90% congruence

between biodiversity hotspots and the global 200 ecore-

gions (for more details see Wikramanayake et al. 2002).

Statements outlined above show evidence of a high-

ly unique and diverse herpetofauna in Sri Lanka. Dur-

ing the last decade Sri Lanka has become recognized as

an amphibian hotspot of high global significance (Mee-

gaskumbura et al. 2002; Pethiyagoda and Manamendra-

Arachchi 1998) and a mega-hotspot of reptile diversity

Figure 6. Lowland rain forest at Sinharaja (top) and montane

forest in the Knuckles Range (bottom; cardamom factory in the

foreground). Photos by Walter R. Erdelen.

(Somaweera and Somaweera 2009). This recognition

may be seen as a bottom-up approach, i.e. a taxon-specif-

ic approach to the issue of prioritizing biodiversity con-

servation, as used in the lUCN lists of threatened fauna

and fiora (see below). It may be seen as an indicator or

a reaction to the fact that overall species and ecosystem

conservation have been biased towards certain taxa (see

above).

The consequence may be use of taxon-specific ap-

proaches to ensure specific characteristics in overall

long-term conservation of species or species analyzed

February 2012 | Volume 5 | Number 2 | e37

amphibian-reptile-conservation.org

042

Erdelen

Figure 7. Variability in geographic distribution among Sri

Lankan reptiles. (A) Chamaeleo zeylanicus, a non-endemic

species of the dry zone lowlands; (B) Naja naja, non-endemic

and found all over the island below some 1500 m asl; (C) Geck-

oella triedrus, a wet zone species which is also locally found

in the dry zone and intermediate zone; (D) Geckoella yakhuna,

restricted to the dry zone lowlands of the north; both species

are endemic to Sri Lanka and need further study as regards to intraspecific variation. The status of the third species occurring in Sri

Lanka (G. collegalensis) is unclear (Somaweera and Somaweera 2009); (E) Rhinophis homolepis, an endemic uropeltid snake found

in the wet zone lowlands; fossorial amphibians and reptiles may be environmental indicators and key groups for an understanding

of species evolution in Sri Lanka (see Cans 1993); (F) Haplocercus ceylonensis, an endemic colubrid snake found in the wet zone

highlands. Photos by Indraneil Das.

(for examples of variation in status and distribution of

species see Figs. 1 and 7). This approach may lead to

a new insight regarding conservation aspects specific to

the herpetofauna of Sri Lanka and be vital for overall or

“holistic” conservation of biodiversity. Concretely, this

approach may relate to rarity, small population sizes, and

patchy geographic distribution of many of Sri Lanka’s

amphibian species.

February 2012 | Volume 5 | Number 2 | e37

amphibian-reptile-conservation.org

043

Sri Lanka’s amphibians and reptiles

lUCN Lists

The 2007 lUCN red list of threatened fauna and flora

of Sri Lanka lists 33% of all vertebrates as nationally

threatened (63% endemic to Sri Lanka). Among major

groups of vertebrates reptiles and amphibians rank first in

numbers of threatened species, followed by bird, mam-

mal, and freshwater fish species (lUCN Sri Lanka and

MoENR 2007).

The 2009 lUCN State of Amphibians of Sri Lanka,

based on a total species number of 105, draws a particu-

larly bleak picture of endangerment: 20% are reported

Extinct, 10% Critically Endangered, 34% Endangered,

6% Vulnerable, and 5% Near-threatened. Only 23% are

of least concern and for 2% insufficient data are avail-

able to assess their status. Sri Lanka ranks highest among

Asian countries, having the greatest percentage of threat-

ened amphibians. It has lost some 20% of its amphib-

ian species during the last century, and over 50% of the

remaining species are prone to extinction (lUCN State

of Amphibians of Sri Lanka, update of 7 April 2009, ac-

cessed through www.iucn.org).

Sri Lanka therefore is not only characterized by the

highest degree of endemism among amphibians in Asia

but also by the highest number of extinct amphibian spe-

cies reported for an individual country. The loss of 20%

of its amphibian species has been a result of human im-

pacts on natural ecosystems during the last 100 years,

particularly to natural forest ecosystems of the wet zone

and central hills of Sri Lanka. It should be noted, how-

ever, that the meaning of “extinct” in this context is not

based on absolute proof but on the lack of more recent

species records.

One hundred and seventy-one indigenous reptile

species, excluding marine species, were assessed by

lUCN (2007). Of these, 16 (9.3%) species are consid-

ered Critically Endangered, 23 (13.5%) Endangered, and

17 (10%) Vulnerable. This translates into a total of 56

(32.7%) species with their existence threatened. Of these,

37 (66%) are species endemic to Sri Eanka.

In the 2007 lUCN list, concern is expressed inter

alia about the facts that: (1) national red lists have not

been integrated into national policies or other ongoing

national conservation actions; (2) better awareness of the

contents of these lists needs to be created among relevant

line ministries; and (3) the status of most threatened spe-

cies has remained unchanged or worsened with time.

These concerns need to be seriously addressed and joint-

ly translated into concrete action by decision makers, the

scientific community, and the public at large.

Institutional arrangement in Sri Lanka

Although this paper focuses on specific issues related to

the conservation of amphibians and reptiles in Sri Lan-

ka, this newer comprehensive understanding presented

needs to be made relevant and tangible within the overall

setup of institutions and agencies managing the environ-

ment, biodiversity, and sustainable development of the

country. The key ministry mandated with sustainable de-

velopment and environmental management in Sri Lanka

is the Ministry of Natural Resources and Environment

(MoENR). MoENR’s regulatory commission is to moni-

tor, revise, and report progress of the Environmental Ac-

tion Plan and to formulate national policies for environ-

mental protection and management. MoENR houses the

National Biodiversity Secretariat who is responsible for

policies and plans for national biodiversity conservation

and attends to national implementation of the Conven-

tion on Biological Diversity (CBD) and the Cartagena

Protocol (see Dela 2009 for further details). The main

sectoral institutions within the MoENR are the Forest

Department, the Department of Wildlife Conservation,

the Central Environmental Authority, and the Marine En-

vironment Protection Authority. An overview of national

stake holders for implementing the CBD and the National

Biodiversity Conservation Action Plan (BCAP) — main

legislation relating to environmental conservation and

management — and key state agencies outside the envi-

ronmental sector dealing with biodiversity conservation

in Sri Lanka are listed in Dela (2009).

De Silva (2001) compiled a list of government de-

partments and organizations which have more specifical-

ly contributed to Sri Lankan herpetology. He lists some

major non-governmental organizations (NGOs) who

specially contribute to improving our knowledge of am-

phibians and reptiles in Sri Lanka. These NGOs are listed

in alphabetic order below (from de Silva 2001; founding

dates are given in brackets where available):

• Amphibia and Reptile Research Organization of Sri

Lanka (ARROS).

• Conservation Breeding Specialist Group (lUCN/

CBSG/SSC), Sri Lanka Network.

• Declining Amphibian Population Task Force, Work-

ing Group Sri Lanka (1999).

• March for Conservation.

• The Neo Synthesis Research Centre.

• The Royal Asiatic Society of Sri Lanka (1 845).

• Snakebite Expert Committee, Sri Eanka Medical

Association (1983).

• Turtle Conservation Project.

• The Wildlife and Nature Protection Society of Sri

Eanka (1894).

• The Wildlife Heritage Trust of Sri Eanka (1990)

• The Young Zoologists Association (1972)

February 2012 | Volume 5 | Number 2 | e37

amphibian-reptile-conservation.org

044

Erdelen

These institutions and agencies have enormous potential

for enhancing efforts to jointly contribute to mainstream-

ing biodiversity conservation into cross-sectoral strate-

gies and plans. This potential applies in particular to the

development aspects and, therefore, for the sustainable

development of Sri Lanka in general. Better cooperation

and planning among conservation stake holders in Sri

Lanka would greatly increase conservation efforts and

are essential in saving the largest portion of biodiversity

in Sri Lanka.

Conservation of Sri Lanka’s

herpetofauna — A proposai

Knowledge of amphibian and reptile geographic distri-

bution in Sri Lanka, especially endemic species, high-

lights the close association between their geographic

distribution patterns and natural ecosystems. For most

species we lack precise information about how species

distributions are linked to specific habitats or microhabi-

tats. This applies in particular to amphibians which show

highly patched distributions and fragmented or small

populations. Further studies are needed to determine if

this is a result of “natural” patchiness, habitat fragmenta-

tion, or sampling artifact (see Janzen and Bopage 2011

for a forest patch herpetofauna study at approximately

1000 m asl).

Studies on extinction risks and population vulner-

ability have not been carried out for most species. Eco-

logical and biogeographical studies are lagging far be-

hind taxonomic and systematic studies. Without doubt,

ecological and biogeographical studies should be con-

tinued and should parallel population studies (including

monitoring of population dynamics), especially in view

of severe habitat fragmentation and additional negative

impacts expected to result from climate change.

All these efforts toward a better understanding of the

status and endangerment of Sri Lanka’s amphibians and

reptiles need not only be sustained but considerably in-

creased. This will require increased support and effort at

national and international levels and must be embedded

in the overall resolve for reinforcing biodiversity conser-

vation in Sri Lanka.

Toward an Action Plan

Many important proposals have been made for the con-

servation of Sri Lanka’s biodiversity and its herpetofauna

(e.g.. Das 1996b; de Silva 2006; lUCN Sri Lanka and

MoENR 2007; Pethyiagoda et al. 2006). These are not

repeated here, but an integrated action plan is proposed

below which focuses on several areas of prime impor-

tance.

1) Mapping existing schemes of cooperation, identi-

fying shortcomings, and providing an optimized

scenario for partnership arrangements at national

and international levels to make the best “use” of

existing capacities.

2) Reinforcing scientific work on the amphibians and

reptiles of Sri Lanka through a targeted approach

and using all national capacities (governmen-

tal institutions and other entities, universities,

NGOs, and other stake holders) and schemes of

international cooperation. Scientific work should

include a continuation of the highly successful

taxonomic work of the past decade but should

increasingly include ecological and biogeo-

graphical work to complement our knowledge of

systematic relationships among taxa (for some

recent problems see Pethyiagoda 2004).

3) Linking this endeavor to work on ecosystem or

plant community classification and conservation

as carried out by Sri Lankan universities, particu-

larly in regards to botanical research or work in

the fields of plant ecology and plant biogeogra-

phy.

4) Developing schemes and scientific programs sup-

ported by the latest space technologies for moni-

toring the status of ecosystems in Sri Lanka for

habitat restoration and recreating continuous hab-

itat or ecosystems (particularly in the wet zone

and central hills). Replanting and reconnecting

forest fragments through planting of indigenous

species, as has been carried out for years by the

Department of Botany at Peradeniya University

(e.g., Ashton et al. 2001).

5) Fostering joint education, research, and degree

work in these fields at universities in Sri Lanka.

This may need to be coordinated among univer-

sities interested in inter-university cooperation.

Such a plan could create better employment op-

portunities and promote qualified staff to work in

conservation and sustainable development sec-

tors.

6) Making biodiversity education more inclusive,

encompassing all levels of the education system

including formal and informal education and ar-

rangements for life-long learning. In addition,

biodiversity education should become part of a

massive effort to champion education for sustain-

able development in the country, closely linked

to public awareness programs, particularly as

needed for the conservation of amphibians and

reptiles.

7) The results of these works should be interconnected

to conservation work carried out by the Sri Lank-

amphibian-reptile-conservation. org 045 February 2012 | Volume 5 | Number 2 | e37

Sri Lanka’s amphibians and reptiles

an government authorities, in particular the For-

est Department, the Department of Wildlife Con-

servation, and the Biodiversity Secretariat.

8) Fostering the role and capacity of the National Mu-

seum in overall conservation efforts for Sri Lank-

an herpetofauna in a national and international

context, and in particular through reinforcing and

facilitating the museum’s international collabora-

tion and programs of work.

9) Reinforcing in situ and ex situ conservation efforts

for amphibians and reptiles in Sri Lanka. The

zoological gardens at Dehiwela and the estab-

lishment of a new facility such as a “Sri Lanka

Aquarium” might generate the needed public at-

tention for the conservation needs of Sri Lanka

and its herpetofauna (see 6).

10) Extending existing activities and programs in na-

tional and international ecotourism programs to

include amphibians and reptiles as specific ex-

amples for creating environmental awareness and

the need for biodiversity conservation.

11) Closer liaison between all stake holders in joint

conservation efforts regarding biodiversity

hotspots of south India’s Western Ghats and Sri

Lanka. A model approach could be developed

for preserving biodiversity in both hotspots

(sometimes considered a single hotspot), serv-

ing as a template for similar analysis in other

biodiversity hotspots. This needs to be based

on a changed mind- set, with a paradigm shift-

ed from “protection” to “conservation,” which

includes active, research-based management

interventions (R. Pethiyagoda, pers. comm.).

For examining the feasibility of such an action plan or a

si mil ar initiative, a workshop or other “kick-off’ meet-

ing with all relevant governmental and non-governmen-

tal stake holders might be a useful first step. A proposed

meeting may contribute to significant positive efforts in

capacity and resource development (a multiple win situ-

ation for all stake holders) and for sustaining Sri Lanka’s

faunal and fioral wealth for future generations.

Conclusions and outlook

Our knowledge of Sri Lanka’s biodiversity has expe-

rienced a quantum leap during the last decade. This is

underscored by massive efforts to scale up taxonomic re-

search, in particular of the fauna of Sri Lanka, which has

led to the discovery of a substantial number of new spe-

cies among invertebrate and vertebrate taxa. Specifically,

genetic studies have contributed to new insights into the

country’s biological diversity. The increase in numbers

of amphibian species scientifically described has been

outstanding, making it the vertebrate group with the

highest percentage of endemic species (some 90%) in Sri

Lanka; also more than twenty new reptile species have

been described during the last decade.

Biodiversity efforts in Sri Lanka need to be further

streamlined between all governmental and non-govern-

mental institutions and agencies. This should include the

consideration of global climate change as possibly the

most important factor affecting the future of Sri Lanka’s

biodiversity, particularly the exceptional biodiversity

in montane areas. A specific focus must be put on con-

nectivity of natural habitat, particularly in the lowland

wet zone and highlands where forests have been severely

fragmented — a phenomenon making these ecosystems

particularly prone to impacts of climate change and ex-

acerbated by the large number of aggressive invasive

alien species now found in the highlands of Sri Lanka (R.

Pethyiagoda, pers. comm.).

The division of institutional activities and the enor-

mous number of ongoing projects related to the conser-

vation of Sri Lanka’s biodiversity may need to be in-

ventoried and mapped at both national and international

levels in order to optimize future efforts. This is espe-

cially needed because of the limited human and finan-

cial resources available to address biodiversity issues

in Sri Lanka. These efforts should be accompanied by

the formation of an inter-institutional coordination plan

for biodiversity research, monitoring, and identification

of threats, as is already proposed in the Fourth Country

Report from Sri Lanka to the Convention on Biological

Diversity (see Dela 2009, Appendix III, p. vii). Such an

initiative may benefit from a regional approach, exchang-

ing experience and addressing common issues especially

since Sri Lanka and the Western Ghats of southern In-

dia are one of the most important global biodiversity

hotspots containing ecoregions of outstanding regional

and global value.

The Decade on Biodiversity (2011-2020) and the

implementation recommendations of the Nagoya COP

10 conference such as the new biodiversity strategy and

the biodiversity targets might offer a unique platform for

launching and sustaining the initiatives outlined here.

This platform could facilitate the release of an updated

National Biodiversity Strategy and Action Plan for Sri

Lanka which might be cast as a living strategic docu-

ment, closely linked to the country’s efforts to imple-

ment sustainable development, with an increased focus

on coping with the effects of global climate change and

using the potential of a green economy.

Acknowledgments. — The manuscript has greatly

benefited from comments by Indraneil Das, Jacques

Richardson, and Rohan Pethyiagoda. “Neil” Das kindly

provided photos, and I am grateful to Rohan for infor-

mation and reference materials. Wilhelm Konle assisted

in the selection of slides. I sincerely thank all these col-

leagues and friends. Special thanks to Craig Hassapakis

February 2012 | Volume 5 | Number 2 | e37

amphibian-reptile-conservation.org

046

Erdelen

who invited me to contribute to the Sri Lanka issue of

Amphibian and Reptile Conservation and assisted in

many ways in publishing the paper. Above all, I would

like to thank my wife Amina, who accompanied me

along the way in writing this paper.

Literature cited

Abeywickrama BA. 1956. The origin and affinities of the fiora

of Ceylon. Proceedings of the IP’' Annual Session of the

Ceylon Association for the Advancement of Science 2:99-

121.

Abeywicbcrama BA. 1993. Integrated conservation for human

survival in Sri Lanka In: Ecology and Landscape Manage-

ment in Sri Lanka. Editors, Erdelen WR, Preu C, Ishwaran

N, Madduma Bandara CM. Margraf Publishers, Weiker-

sheim, Germany. 15-24.

AmarasingheAAT, T iedemannF, KarunarathnaDMS S . 20 1 1 .

Calotes nigrilabris Peters, 1860 (Reptilia: Agamidae; Dra-

coninae): A threatened highland agamid lizard in Sri Eanka.

Amphibian and Reptile Conservation 5(2):33-43(e32).

Ariyasiri K, Bowatte G, Menike U, Meegasukumbura S,

Meegasukumbura M. 2011. Predator-induced plasticity in

tadpoles of Polypedates cruciger (Anura: Rhacophoridae).

Amphibian and Reptile Conservation 5(2): 14-21(e29).

Ashton MS, Gunatilleke CVS, Singhakumara BMP, Guna-

TiLLEKE lAUN. 2001. Restoration pathways for rain forest

in southwest Sri Lanka: A review of concepts and models.

Forest Ecology and Management 154(3):409-430.

Ashton PS, Gunatilleke CVS. 1987. New light on the plant

geography of Ceylon. I. Historical plant geography. Journal

of Bio geography 14(3);249-285.

Becker CG, Fonseca CR, Haddad CEB, Batista RF, Prado PI.

2007. Habitat split and the global decline of amphibians.

Nature 318:1775-1777.

Bahir mm, Surasinghe TD. 2005. A conservation assessment

of the Sri Eankan Agamidae (Reptilia: Sauria). The Raffles

Bulletin of Zoology, Supplement 12:407-412.

Beenaerts N, Pethyiagoda R, Ng PKE, Yeo DJC, Bex GJ,

Bahir MM, Artois T. 2010. Phylogenetic diversity of Sri

Lankan freshwater crabs and its implications for conserva-

tion. Molecular Ecology 19(1): 183- 196.

Bhimachar BS. 1945. Zoogeographical divisions of the West-

ern Ghats as evidenced by the distribution of hill stream

fishes. Current Science 14:12-16.

Biswas S. 2008. Did biotic impoverishment facilitate phe-

nomenal diversification in Sri Lanka? Current Science

95(8):1021-1025.

Biswas S, Pawar SS. 2006. Phylogenetic tests of distribution

patterns in South Asia: Towards an integrative approach.

Journal of Bioscience 31(1):95-113.

Blaneord WT. 1870. Notes on some Reptilia and Amphibia

from Central India. Journal of the Asiatic Society of Bengal

39:335-376.

Boulenger GA. 1890. The Fauna of British India, including

Ceylon and Burma. Reptilia and Batrachia. Taylor and

Francis, London, UK.

Cadle JE, Dessauer HC, Gans C, Gartside DF. 1990. Phylo-

genetic relationships and molecular evolution in uropeltid

snakes (Serpentes: Uropeltidae): Allozymes and albumin

immunology. Biological Journal of the Linnean Society

40(3):293-320.

CiNCOTTA RP, WiSNEWSKi J, Engelman R. 2000. Human popula-

tion in the biodiversity hotspots. Nature 404:990-992.

Clarke CB. 1898. Subareas of British India illustrated by the

detailed distribution of the Cyperaceae in that Empire. Jour-

nal of the Linnean Society London 34: 1-146.

CooRAY PG. 1984. An Introduction to the Geology of Sri Lanka

( Ceylon). Government Press, Colombo, Sri Lanka.

Crusz H. 1973. Nature conservation in Sri Lanka (Ceylon).

Biological Conservation 5(3): 199-208.

Crusz H. 1984. Parasites of endemic and relict vertebrates:

A biogeographical review In: Ecology and Biogeography

in Sri Lanka. Editor, Fernando CH. Junk Publishers, The

Hague, The Netherlands. 321-351.

Crusz H. 1986. The vertebrates of Sri Lanka: Endemism and

other aspects. Report of the Society for Research in Native

Livestock (Japan) 11:65-80.

Crusz H, Nugaliyadde L. 1978. Parasites of the relict fauna of

Ceylon. VIE General considerations and first host-parasite

checklist. Compte rendu des seances de la societe de bio-

geographie 477:85-106.

Das I. 1996a. Biogeography of the Reptiles of South Asia.

Krieger Publishing Company, Malabar, Florida, USA.

Das I. 1996b. The South Asian Herpetofauna. A Conservation

Action Plan. Compiled by Indraneil Das and Members of

the lUCN/SSC South Asian Reptile and Amphibian Special-

ist Group. Unpublished.

Das I, DE Silva A, Austin CC. 2008. Anew species of Eutropis

(Squamata: Scincidae) from Sri Lanka. Zootaxa 1700:35-

52.

DeAlwis L. 1969. The National Parks of Ceylon. A guide. Gov-

ernment Press, Colombo, Sri Lanka.

Dela JDS. 2009. Fourth Country Report from Sri Lanka to the

United Nations Convention on Biological Diversity. Co-

lombo, Sri Lanka.

Deraniyagala pep. 1939. The Tetrapod Reptiles of Ceylon -

Testudinates and Crocodilians. Volume 1. Government

Press, Colombo, Sri Lanka.

Deraniyagala PEP. 1953. A Colored Atlas of some Vertebrates

from Ceylon. Volume II. Tetrapod Reptilia. Government

Press, Colombo, Sri Lanka.

Deraniyagala PEP. 1955. A Colored Atlas of some Vertebrates

from Ceylon. Volume III. Serpentoid Reptilia. Government

Press, Colombo, Sri Lanka.

Deraniyagala SU. 1993. Pleistocene human ecology in Sri

Lanka In: Ecology and Landscape Management in Sri Lan-

ka. Editors, Erdelen WR, Preu C, Ishwaran N, Madduma

Bandara CM. Margraf Publishers, Weikersheim, Germany.

25-34.

DE Silva A. 1990. Colour Guide to the Snakes of Sri Lanka. R &

A Publishing Ltd., Avon, UK.

DE Silva A. 1994. The amphibia of Sri Lanka: A provisional

February 2012 | Volume 5 | Number 2 | e37

amphibian-reptile-conservation.org

047

Sri Lanka’s amphibians and reptiles

checklist and their common names. Lyriocephalus 1:20-26.

DE Silva A. 1996. The Amphibia of Sri Lanka: A Checklist and

an Annotated Bibliography. Department of Wildife Conser-

vation/GEF/UNDP/FAO, Kandy, Sri Lanka.

DE Silva A. 1998a. The Sauria (Lizards and Varanids) of Sri

Lanka: A Checklist and an Annotated Bibliography. Depart-

ment of Wildife Conservation/GEF/UNDP/FAO, Kandy, Sri

Lanka.

DE Silva A. 1998b. The Snakes of Sri Lanka: A Checklist and an

Annotated Bibliography. Department of Wildife Conserva-

tion/GEF/UNDP/FAO, Kandy, Sri Lanka.

DE Silva A. 1998c. The Testudines and Crocodilians of Sri

Lanka: A Checklist and an Annotated Bibliography. Depart-

ment of Wildlife Conservation/GEF/UNDP/FAO, Kandy,

Sri Lanka.

DE Silva A. (Editor). 1998. Biology and Conservation of the

Amphibians, Reptiles and their Habitats in South Asia. Pro-

ceedings of the International Conference on the Biology

and Conservation of the Amphibians and Reptiles of South

Asia, held at the Institute of Fundamental Studies. Kandy

and University of Peradeniya, Sri Lanka, August 1-5, 1996,

Amphibia and Reptile Research Organization of Sri Lanka,

Peradeniya, Sri Lanka.

DE Silva A. 2001. The Herpetofauna of Sri Lanka: An Histori-

cal Overview, Current Status with Checklists. Amphibia and

Reptile Research Organization of Sri Lanka, Peradeniya, Sri

Lanka.

DE Silva A. 2006. Current status of the reptiles of Sri Lanka

In: The Fauna of Sri Lanka: Status of Taxonomy, Research

and Conservation. Editor, Bambaradeniya CNB. lUCN Sri

Lanka, Colombo, Sri Lanka. 134-163.

DE Silva PHDH. 1955. A list of Amphibia recorded from Cey-

lon with a report on the Amphibia in the Colombo Museum.

Spolia Zeylanica 27:244-250.

DE Silva PHDH. 1977. Colombo Museum 100 Years 1877-1977

Souvenir. Lake House Publishing, Colombo, Sri Lanka.

DE Silva PHDH. 1980. Snake Fauna of Sri Lanka with Special

Reference to Skull, Dentition and Venom in Snakes. Govern-

ment Press, Colombo, Sri Lanka.

Dietz RS, Holden JC. 1970. The breakup of Pangea. Scientific

American 223:30-41.

Dodd CK, Bartholomew B. 2002. Fourth World Congress of

Herpetology: 3-9 December 2001, Bentota, Sri Lanka. Her-

petological Review 33(l):3-5.

Dutta SK. 1985. Amphibians of India and Sri Lanka. Ph.D.

Dissertation, University of Kansas. Unpublished.

Dutta SK, Manamendra-Arachchi K. 1996. The Amphibian

Fauna of Sri Lanka. Wildlife Heritage Trust of Sri Lanka,

Colombo, Sri Lanka.

Erdelen WR. 1977. Intraspezifische Variabilitdt bei Calotes

versicolor Daudin (Sauria: Agamidae). Diploma Thesis,

University of Munich, Germany. Unpublished.

Erdelen WR. 1983. Die Gattung Calotes in Sri Lanka: Bezie-

hungen innerhalb eines Artensystems tropischer Echsen.

Dissertation Thesis, University of Munich, Germany. Un-

published.

Erdelen WR. 1984. The genus Calotes (Sauria, Agamidae) in

Sri Lanka: Distribution patterns. Journal of Bio geography

ll(6):515-525.

Erdelen WR. 1988a. Population dynamics and dispersal in

three species of agamid lizards in Sri Lanka: Calotes calo-

tes, C. versicolor and C. nigrilabris. Journal of Herpetology

22(l):42-52.

Erdelen WR. 1988b. Forest ecosystems and nature conserva-

tion in Sri Lanka. Biological Conservation 43(2): 115-135.

Erdelen WR. 1989. Aspects of the biogeography of Sri Lanka

In: Forschungen auf Ceylon 111. Editor, Schweinfurth U.

Steiner Verlag, Stuttgart, Germany. 73-100.

Erdelen WR. 1993a. Biogeographische Muster und Prozesse.

Die tropische Kontinentalinsel Sri Lanka. Habilitations-

schrift. University of the Saarland, Saarbrucken, Germany.

Unpublished.

Erdelen WR. 1993b. The relevance of scale and hierarchy con-

cepts for landscape management. A Sri Lankan perspective

In: Ecology and Landscape Management in Sri Lanka. Edi-

tors, Erdelen WR, Preu C, Ishwaran N, Madduma Bandara

CM. Margraf Publishers, Weikersheim, Germany. 221-224.

Erdelen WR. 1996. Tropical rain forests in Sri Lanka: Char-

acteristics, history of human impact, and the protected area

system In: Tropical Rainforest Research - Current Issues.

Editors, Edwards DS, Booth WE, Choy SC. Kluwer Aca-

demic Publishers, Dordrecht, The Netherlands. 503-511.

Erdelen WR, Preu C. 1990a. Quaternary coastal and vegeta-

tion dynamics in the Palk Strait region. South Asia - The

evidence and hypotheses In: Vegetation and Erosion. Editor,

Thornes JB. John Wiley & Sons, Chichester, UK. 491-504.

Erdelen WR, Preu C. 1990b. Background and outcome of an

international symposium on “Ecology and Landscape Man-

agement in Sri Lanka.” GeoJournal 22(2):215-217.

Erdelen WR, Preu C, Ishwaran N, Madduma Bandara CM.

Editors. 1993. Ecology and Landscape Management in Sri

Lanka. Margraf Publishers, Weikersheim, Germany.

Ferguson W. 1877. Reptile Fauna of Ceylon. Letter on a col-

lection sent to the Colombo Museum. Government Printers,

Colombo, Sri Lanka.

Fernando P, Dayawansa N, Siriwardena M. 1994. Bufo ko-

tagamai, a new toad (Bufonidae) from Sri Lanka. Journal

of South Asian natural History 1 ( 1 ): 1 1 9- 1 24.

Fernando SS, Wickramasingha LJM, Rodirigo RK. 2007. A

new species of endemic frog belonging to genus Nannoph-

rys Gunther, 1869 (Anura: Dicroglossinae) from Sri Lanka.

Zootaxa 1403:55-68.

Gans C. 1993. Fossorial amphibians and reptiles: Their distri-

butions as environmental indicators In: Ecology and Land-

scape Management in Sri Lanka. Editors, Erdelen WR, Preu

C, Ishwaran N, Madduma Bandara CM. Margraf Publishers,

Weikersheim, Germany. 189-199.

Gower DJ, Maduwage K. 20 1 1 . Two new species of Rhinophis

Hemprich (Serpentes: Uropeltidae) from Sri Lanka. Zoo-

taxa 2881:51-68.

Gunatilleke CVS, Gunatilleke IAUN. 1983. The floristic

composition of Sinharaja - a rain forest in Sri Lanka with

special reference to endemics. The Sri Lanka Forester 14(3-

4): 170-179.

February 2012 | Volume 5 | Number 2 | e37

amphibian-reptile-conservation.org

048

Erdelen

Gunatilleke IAUN, Gunatilleke CVS. 1984. Distribution of

endemics in the tree flora of a lowland hill forest in Sri Lan-

ka. Biological Conservation 28(3):275-285.

Gunatilleke N, Pethyiagoda R, Gunatilleke S. 2008. Biodi-

versity of Sri Lanka. Journal of the National Science Foun-

dation 36 (Special Issue):23-59.

Gunther ACLG. 1858. On the geographical distribution of

reptiles. Proceedings of the Zoological Society of London

21(l):373-398.

Gunther ACLG. 1864. The Reptiles of British India. London,

UK.

Haly a. 1886a. First Report on the Collection of Batrachia in

the Colombo Museum. Colombo, Sri Lanka.

Haly A. 1886b. First Report on the Collection of Snakes in the

Colombo Museum. Colombo, Sri Lanka.

Haly A. 1891. Report on the Collection ofReptilia and Batra-

chia in the Colombo Museum. Government Press, Colombo,

Sri Lanka.

Hooker JD. 1906. A Sketch of the Flora of British India. Lon-

don, UK.

Hooker JD, Thomson T. 1855. Flora Indica. W. Pamplin, Lon-

don, UK.

IPCC 2007. Climate Change 2007. Fourth Assessment Report

of the Intergovernmental Panel on Climate Change. Cam-

bridge University Press, New York, USA.

IskandarDT, Erdelen WR. 2006. Conservation of amphibians

and reptiles in Indonesia: Issues and problems. Amphibian

and Reptile Conservation 4(l);60-87.

lUCN 2009. State of Amphibians of Sri Lanka, update of 7

April 2009. [Online]. Available: www.iucn.org [Accessed:

20 October 2011].

lUCN Sri Lanka and MoENR (Ministry oe Environment and

Natural Resources). 2007. The 2007 Red List of Threat-

ened Fauna and Flora of Sri Lanka. Colombo, Sri Lanka.

Janzen P, Bopage M. 2011. The herpetofauna of a small and

unprotected patch of tropical rainforest in Momingside,

Sri Lanka. Amphibian and Reptile Conservation 5(2): 1-

13(e26).

Jayasuriya AHM. 1984. Flora of Ritigala Natural Reserve. The

Sri Lanka Forester 16:61-156.

Jayasuriya AHM, Pemadasa MA. 1983. Factors affecting the

distribution of tree species in a dry zone montane forest in

Sri Lanka. Journal of Ecology 71(2):571-583.

Jerdon TC. 1 862- 1 864. The Birds of India. Published by author.

Calcutta, India.

Katz MB. 1978. Sri Lanka in Gondwanaland and the evolution

of the Indian Ocean. Geological Magazine 115:237-244.

Keast a. 1973. Contemporary biotas and the separation se-

quence of the southern continents In: Implications of Con-

tinental Drift to the Earth Sciences. Volume 1 . Editors, Tar-

ling DH, Runcorn SK. Academic Press, New York. 309-343.

Kelaart EF. 1852. Ceylon Amphibia. Prodromus Faunae Zey-

lanicae 1:191-197.

Kirtisinghe P. 1957. The Amphibia of Ceylon. Published by the

author. Colombo, Sri Lanka.

Klaus S, Schubart CD, Streit B, Peenninger M. 2010. When

Indian crabs were not yet Asian - biogeographic evidence

amphibian-reptile-conservation.org

for Eocene proximity of India and Southeast Asia. BMC

Evolutionary Biology 10:287.

Kotagama SW, Arudpragasam KD, Kotalawala I. 1981.

A Check List of Amphibia of Sri Lanka. UNESCO: MAB

Publication 10. National Science Council of Sri Lanka, Co-

lombo, Sri Lanka.

Kuper W, Sommer JH, Lovett JC, Mur keJ, Linder HP, Beentje

HJ, VAN Rompaey RASR, Chatelain C, Sosee M, B arthlott

W. 2004. African hotspots of biodiversity redefined. Annals

of the Missouri Botanical Garden 91(4):525-535.

Ladle RJ, Whittaker RJ. (Editors). 2011. Conservation Bioge-

ography. John Wiley & Sons Etd, Chichester, UK.

MaduwageK,SilvaA,Manamendra-ArachchiK, Pethyiago-

da R. 2009. A taxonomic revision of the South Asian hump-

nosed pit vipers (Squamata: Viperidae: Hypnale). Zootaxa

2232:1-28.

Manamendra-Arachchi K, Gabadage D. 1996. Limnonectes

kirtisinghei, a new species of ranid frog from Sri Lanka.

Journal of South Asian natural History 2:31-42.

Mani ms. Editor. 1974. Ecology and Biogeography in India.

Junk Publishers, The Hague, The Netherlands.

Marris E. 2009. Ragamuffin Earth. Nature 460:450-453.

Maxson ER. 1984. Albumin and Australian frogs: Mo-

lecular data a challenge to speciation model. Science

225(4665):957-958.

McKenna MC. 1975. Fossil mammals and early Eocene North

Atlantic continuity. Annals of the Missouri Botanical Gar-

den 62(2):335-353.

ME A (Millennium Ecosystem Assessment). 2005. Ecosystems

and Human Well-being: Biodiversity Synthesis. World Re-

sources Institute, Washington, DC, USA.

Meegaskumbura M, Bossuyt F, Pethyiagoda R, Manamen-

dra-Arachchi K, Bahir M, Milinkovitch MC, Schnei-

der CJ. 2002. Sri Lanka: An amphibian hotspot. Science

298(5592):379.

Ministry oe Forestry and Environment. 1999. Biodiversity

Conservation in Sri Lanka: A Framework for Action. Min-

istry of Forestry and Environment, Battaramulla, Sri Lanka.

Mutke J, B arthlott W. 2005. Patterns of vascular plant di-

versity at continental to global scales. Biologiske Skrifter

55:521-537.

Mutke J, Sommer JH, Kreet H, Kier G, B arthlott W. 2011.

Vascular plant diversity in a changing world: Global cen-

tres and biome-specific patterns In: Biodiversity Hotspots.

Editors, Zachos FE, Habel JC. Springer- Verlag, Berlin, Ger-

many. 83-96.

Myers N. 1988. Threatened biotas: “hot spots” in tropical for-

ests. The Environmentalist 8(3): 187-208.

Myers N. 1990. The biodiversity challenge: Expanded hot-

spots analysis. The Environmentalist 10(4):243-256.

Myers N, Mittermeier RA, Mittermeier CG, da Fonseca GAB,

Kent J. 2000. Biodiversity hotspots for conservation priori-

ties. Nature 365:853-858.

Oliver PM, Adams M, Fee MS Y, Hutchinson MN, Doughty P.

2009. Cryptic diversity in vertebrates: Molecular data dou-

ble estimates of species diversity in a radiation of Australian

lizards {Diplodactylus, Gekkota). Proceedings of the Royal

February 2012 | Volume 5 | Number 2 | e37

049

Sri Lanka’s amphibians and reptiles

Society B 21 6-2QQ\-2QQl.

Olson DM, Dinerstein E. 1998. The Global 200: A represen-

tation approach to conserving the earth’s distinctive ecore-

gions. Conservation Biology 12(3):502-515.

Olson DM, Dinerstein E, Wikramanayake ED, Burgess ND,

Powell GVN, Underwood EC, D’Amico JA, Itoua I,

Strand HE, Morrison JC, Loucks CJ, Allnutt TF, Rick-

etts TH, Kura Y, Lamoreux JF, Wettengel WW, Hedao

P, Kassem KR. 2001. Terrestrial ecoregions of the world: A

new map of life on Earth. BioScience 51(ll):933-938.

Pearson DL, Ghorpade K. 1 989. Geographical distribution and

ecological history of tiger beetles (Coleoptera: Cicindeli-

dae) of the Indian subcontinent. Journal of Biogeography

16(4):333-344.

Pethiyagoda R. 2004. Biodiversity law has had some unintend-

ed effects. Nature 429; 129.

Pethiyagoda R. 2007. Pearls, Spices and Green Gold. An Il-

lustrated History of Biodiversity Exploration in Sri Lanka.

World Heritage Trust Publications, Colombo, Sri Lanka.

Pethiyagoda R, Manamendra- Arachchi K. 1998. Evaluating

Sri Lanka’s amphibian diversity. Occassional Papers of the

Wildlife Heritage Trust 2:1-11.

Pethiyagoda R, Manamendra- Arachchi K, Bahir MM, Mee-

GASKUMBURA M. 2006. Sri Lankan amphibians: Diversity,

uniqueness and conservation In: The Fauna of Sri Lanka:

Status of Taxonomy, Research and Conservation. Editor,

Bambaradeniya CNB. lUCN Sri Lanka. Colombo, Sri Lan-

ka. 125-133.

Phillips WWA. 1942. The distribution of mammals in Ceylon

- with special reference to the need for wild life sanctuaries.

Loris 2:4-9.

PiELOU EC. 1979. Biogeography. John Wiley & Sons, New

York, New York, USA.

Prain D.1903. Bengal Plants I. Botanical Survey of India. Cal-

cutta, India.

Premathilake R, Risberg J. 2003. Late Quaternary climate his-

tory of the Horton Plains, Sri Lanka. Quaternary Science

Reviews 22(14); 1525- 1541.

Prendergast JR, Quinn RM, Lawton JH, Eversham SC, Gib-

bons DW. 1993. Rare species, the coincidence of diversity

hotspots and conservation strategies. Nature 365:335-337.

Raheem DC, Naggs F, Chimonides PDJ, Preece RC, Eggleton

P. 2009. Eragmentation and pre-existing species turnover

determine land-snail assemblages of tropical rain forest.

Journal of Bio geography 36(10): 1923-1938.

Raven PH, Axelrod DJ. 1974. Angiosperm biogeography and

past continental movements. Annals of the Missouri Botani-

cal Garden 61(3);539-673.

Reid WV. 1998. Biodiversity hotspots. Trends in Ecology and

Evolution 13(7):275-280.

Roberts JD, Maxson LR. 1985a. Tertiary speciation models in

Australian Anurans: Molecular data challenge Pleistocene

scenario. Evolution 39(2):325-334.

Roberts JD, Maxson LR. 1985b. The biogeography of south-

ern Australian frogs: Molecular data reject multiple inva-

sion and Pleistocene divergence models In: Biology of

Australasian Frogs and Reptiles. Editors, Grigg G, Shine

R, Ehrmann H. Surrey Beatty and Sons, Chipping Norton,

Australia. 83-89.

SCBD (Secretariat oe the Convention on Biological Di-

versity). 2010. Global Biodiversity Outlook 3. Montreal,

Canada.

Schulte II JA, Macey JR, Pethiyagoda R, Larson A. 2002.

Rostral horn evolution among Agamid lizards of the genus

Ceratophora endemic to Sri Lanka. Molecular Phylogenet-

ics and Evolution 22(1);111-117.

Smith MA. 1931. The Fauna of British India, including Ceylon

and Burma. Reptilia and Amphibia. Volume I. - Loricata

and Testudines. Taylor and Francis, London, UK.

Smith MA. 1935. The Fauna of British India, including Cey-

lon and Burma. Reptilia and Amphibia. Volume II. - Sauria.

Taylor and Francis, London, UK.

Smith MA.1943. The Fauna of British India, including Ceylon

and Burma. Reptilia and Amphibia. Volume III. - Serpentes.

Taylor and Francis, London, UK.

SoMASEKARAM T. (Editor) 1988. The National Atlas of Sri Lan-

ka. Survey Department, Colombo, Sri Lanka.

SoMAWEERA R, SoMAWEERA N. 2009. Lizards of Sri Lanka. A

Colour Guide with Field Keys. Edition Chimaira, Erankfurt

am Main, Germany.

Stern N. 2006. Stern Review: The Economics of Climate

Change. HM Treasury, London, UK.

Stuart BL, Inger RF, Voris HK. 2006. High level of cryptic

species diversity revealed by sympatric lineages of South-

east Asian forest frogs. Biology Letters 2:470-474.

Swan B. 1983. An Introduction to the Coastal Geomorphology

of Sri Lanka. Government Press, Colombo, Sri Lanka.

Taylor EH. 1947. Comments on Ceylonese snakes of the ge-

nus Typhlops with descriptions of new species. University of

Kansas Science Bulletin 31(13):283-298.

Taylor EH. 1950a. Ceylonese lizards of the family Scincidae.

University of Kansas Science Bulletin 33(13):481-518.

Taylor EH. 1950b. A brief review of Ceylonese snakes. Uni-

versity of Kansas Science Bulletin 33(14):519-601.

Taylor EH. 1953a. A review of the lizards of Ceylon. Univer-

sity of Kansas Science Bulletin 35(1 2) :15 25 -1585.

Taylor EH. 1953b. A report on a collection of Ceylonese ser-

pents. University of Kansas Science Bulletin 35(14): 1615-

1624.

TEEB 2010. The Economics of Ecosystems and Biodiversity:

Ecological and Economic Foundations. Editor, Kumar P.

Earthscan, London, UK.

ViEITES DR, WOLLENBERG KC, AnDREONE F, KoHLER J, GlAW

F, Vences M. 2009. Vast underestimation of Madagascar’s

biodiversity evidenced by an integrative amphibian inven-

tory. Proceedings of the National Academy of Sciences of

the Unites States of America 106(20):8267-8272.

Wadia DN. 1976. Geology of India. Tata McCraw-Hill Pub-

lishers Company, New Delhi, India.

Wait CCS. 1914. The distribution of birds in Ceylon and its

relation to recent geological changes in the island. Spolia

Zeylanica 10:1-32.

Wikramanayake E, Dinerstein E, Loucks CJ, OlsonDM, Mor-

rison J, Lamoreux J, McKnight M, Hedao P. 2002. Terres-

February 2012 | Volume 5 | Number 2 | e37

amphibian-reptile-conservation.org

050

Erdelen

trial E coregions of the Indo-Pacific. A Conservation Assess-

ment. Island Press, Washington, D.C., USA.

Willis JC. 1916. The evolution of species in Ceylon, with refer-

ence to the dying out of species. Annals of Botany 30:1-23.

Manuscript received: 31 October 2011

Accepted: 14 November 2011

Published: 29 February 2012

WALTER R. ERDELEN studied zoology, botany, genetics,

and chemistry and obtained a doctorate in ecology and zoology

from the University of Munich, Germany, where he worked

as lecturer and researcher. He held professorship positions in

ecology and biogeography at German universities and at the

Bandung Institute of Technology, Indonesia. He carried out ex-

tensive research in the areas of biodiversity, ecology, biogeog-

raphy, conservation biology, evolutionary biology, animal mor-

phology and systematics, with a strong focus on herpetology.

He worked in the tropics of south and southeast Asia and

Africa, including several years of ecological and herpetologi-

cal field research and university cooperation and education

programs in Sri Lanka, India, and Indonesia. He has published

books and numerous scientific papers in his fields of specializa-

tion. He is affiliated with many national and international pro-

fessional associations and worked as a consultant for national

and international organizations, where he has been entrusted

with advisory and evaluation tasks especially related to ecolog-

ical research, biodiversity conservation and capacity building

programs in developing countries.

As Assistant Director-General for Natural Sciences of the

United Nations Educational, Scientific and Cultural Organiza-

tion (UNESCO) he had been responsible for the Organization’s

programs and activities in the area of natural and ecological

sciences. He is based near Paris, Prance.

February 2012 | Volume 5 | Number 2 | e37

amphibian-reptile-conservation.org

051

Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium,

provided the original author and source are credited.

Amphibian & Reptiie Conservation 5(2):52-64.

Range extension for Duttaphrynus /cofagama/ (Amphibia:

Bufonidae) and a preliminary checklist of herpetofauna from

the Uda Maliboda Trail in Samanala Nature Reserve, Sri Lanka

^ ^INDIKA PEABOTUWAGE, ^'■1. NUWAN BANDARA, *^DINAL SAMARASINGHE, ^NIRMALA PERERA,

^MAJINTHA MADAWALA, ^‘'CHAMARA AMARASINGHE, K. DUSHANTHA KANDAMBI, AND

M. S. SURANJAN KARUNARATHNA

^Department of Botany, Faculty of Science, University of Peradeniy a, Peradeniya, SRI LANKA ^Youth Exploration Society of Sri Lanka, PO Box

82, Gannoruwa, SRI LANKA ^Young Zoologists’ Association, Department of National Zoological Gardens, Dehiwala 10350, SRI LANKA '^“El-

langaawa” Unity care for Community & Nature, No: 1/112, Hapugoda, Ambatenna 20136, SRI LANKA ^Nature Exploration & Education Team,

B-l/G-6, De Soysapura Elats, Moratuwa 10400, SRI LANKA

Abstract. — Uda Maliboda Trail is an unstudied, remarkable forest located in the northwest region

of Samanala Nature Reserve (SNR) in Sri Lanka’s wet zone. Here we report the first record of D.

kotagamai from Uda Maliboda Trail and the lowest elevation records of four highland Rhacophorid

frogs: Pseudophiiautus aito, R asankai, P. femoral is, and Taruga eques. Further, we present results

of a preliminary study of herpetofaunal diversity in Uda Maliboda Trail. Thirty-four amphibian (26

endemic and 19 Threatened) and 59 reptile (32 endemic and 19 Threatened) species were observed.

This wet zone forest supports high herpetofaunal diversity; however activities such as deforesta-

tion, human encroachment, mining, agriculture, dumping, road construction, and a hydroelectric

power station threaten the ecology of this biologically diverse forest.

Key words. Amphibians, awareness, conservation, Duttaphrynus, global biodiversity hotspot, Pseudophiiautus,

reptiles, Sri Lanka, threatened, wet zone

Citation: Peabotuwage I, Bandara IN, Samarasinghe D, Perera N, Madawala M, Amarasinghe C, Kandambi HKD, Karunarathna DMSS. 2012. Range

extension for Duttaphrynus kotagamai (Amphibia: Bufonidae) and a preliminary checklist of herpetofauna from the Uda Maliboda Trail in Samanala

Nature Reserve, Sri Lanka. Amphibian & Reptile Conservation 5(2):52-64 (e38).

Introduction

Western Ghats and Sri Lanka have collectively been des-

ignated a global biodiversity hotspot (Mittermeier et al.

2004; Myers et al. 2000). Favorable environmental fac-

tors such as high rainfall, humidity, and a high density

of undergrowth vegetation in this region have assisted in

sustaining regional diversity and distinctness (Bossuyt et

al. 2005; Gunawardene et al. 2007). Sri Lanka comprises

the smaller portion of the hotspot, with a total land area

of 65,610 km^. Despite its small size, the region has a

spectacular assemblage of amphibians and reptiles. Re-

cent molecular studies on amphibians (Rhacophorids

and Caecilians) and Uropeltid snakes have shown that

Sri Lanka has maintained a fauna distinct from the In-

dian mainland (Bossuyt et al. 2004; Meegaskumbura et

al. 2002; Pethiyagoda 2005), yet these subregions are

separated only by about 300 kilometers (direct distance).

Of Sri Lanka’s three major climatic zones (wet, in-

termediate, and dry) the wet zone harbors a significant-

ly high level of herpetofaunal diversity and endemism

(Bambaradeniya et al. 2003; Senanayake et al. 1977;

Wijesinghe and Dayawansa 2002). The wet zone receives

abundant rainfall (annual average 3,000 mm), has con-

siderable forest cover, and maintains favorable humid-

ity and temperatures to support such high herpetofaunal

diversity. Previous studies have noted that some herpeto-

faunal species as well as the wet zone forests themselves

are threatened due to a variety of human activities (e.g.,

lUCN-SL and MENR-SL 2007). Many wet zone forests

have yet to be studied. Uda Maliboda in the Kegalle dis-

trict (Sabaragamuwa Province) is one such unstudied wet

zone forest.

Kotagama’s dwarf toad {Duttaphrynus kotagamai)

is endemic and Endangered and is one of the rarest

bufonids in Sri Lanka (De Silva 2009). Originally de-

scribed from the Sinharaja World Heritage Site in 1994

by Prithiviraj Fernando and Nihal Dayawansa (Fernando

et al. 1994) this toad is known only from the Kitulgala,

Massena, Erathna, and Delwala forest areas (Dutta and

Manamendra-Arachchi 1996; Goonatilake and Goonati-

Correspondence. Email: ^ dmsameera@gmail.com and *dinal.salvator@ gmail.com

January 2012 | Volume 5 | Number 2 | e38

amphibian-reptile-conservation.org

052

Peabotuwage et al.

lake 2001). It favors a few primary lowland rain forests

in the wet zone with elevations below 1,070 m (lUCN-

SL 2011). According to Manamendra-Arachchi and

Pethiyagoda (2006) the holophoront (USNM 311595 H)

has been lost from the National Museum of Natural His-

tory, Washington, D.C. (USA). Herein we describe new

localities and a range extension for D. kotagamai from

a lowland rain forest in the northwestern boundary of

the Samanala Nature Reserve (SNR) and further provide

a preliminary checklist of herpetofauna from the Uda

Maliboda Forest area.

Materials and methods

We used visual encounter survey methods (Crump and

Scott 1994) to conduct herpetofaunal surveys for a to-

tal of 17 days and nights between 2006 and 2011. Night

searches were performed using headlamps and flash-

lights. We searched specific microhabitats including un-

derneath stones and decaying logs, inside tree holes, and

other potential herpetofaunal retreats. Road kills and data

from animals dispatched by villagers were also used as

sources of information. Specimens were hand captured,

photographed, identified using held guides and scientific

publications (Ashton et al. 1997; De Silva 2009; Dutta

and Manamendra-Arachchi 1996; Maduwage et al 2009;

Manamendra-Arachchi et al. 2007; Manamendra-Arach-

chi and Pethiyagoda 2006; Meegaskumbura et al. 2010;

Somaweera 2006; Somaweera and Somaweera 2009; Vo-

gel and Rooijen 2011; Wickramasinghe et al. 2007a, b).

and then released back to the original capture site without

injury. Species nomenclature was based on Frost et al.

(2006), Kotaki et al. (2010), Sumida et al. (2007), and

Senaratna (2001), and conservation status was evaluated

on the lUCN-SL and MENR-SL (2007).

Study area and habitats

The Samanala Nature Reserve (SNR) is one of the larg-

est and most important forest areas for endemic biodiver-

sity in Sri Lanka and is owned by the Central Highlands

World Heritage Centre (UNESCO 2011). The Study area

lies between 6°53’01.58” N and 80°26’31.18” E with

elevations ranging from 300-700 m (Fig. 1). This forest

area is part of the Kegalle district in Sabaragamuwa Prov-

ince. Average annual rainfall ranges from 3,000-4,500

mm and the average annual temperature is 27.9 °C (Fig.

2). The vegetation of Uda Maliboda Trail is categorized

as lowland wet evergreen forest (Gunatilleke and Guna-

tilleke 1990) and is comprised of the following dominant

genera: Doona, Stemonoporus, Calophyllum, Syzygium,

Shorea, Dipterocarpus, Cullenia, and Mesua (Table 1).

Pilgrims use four main trails annually between Decem-

ber and April to reach Adams Peak to worship. The Uda

Maliboda Trail starts from the “Uda Maliboda village”

and continues through Madahinna (Kuruwita trail) via

Adams Peak (elevation 2,245 m). This is the longest trail

and is seldom used by pilgrims since it consists of rough

terrain and narrow foot paths (Karunarathna et al. 2011).

LKm

. * -7“ p.Eraihna

dUda Maliboda

T+fummodara

Figure 1. Map of study area (sky view source: Google map).

January 2012 | Volume 5 | Number 2 | e38

amphibian-reptile-conservation.org

053

Uda Maliboda trail and a preliminary herpetofaunal checklist

Table 1. Floral species presence in different level of Uda Maliboda area (Uda Maliboda Trail in SNR).

Prominent layer

Plant species diversity

Canopy

Adinandra lasiopetala, Bhesa ceylanica, Calophyllum trapezifolium, Cullenia ceylanica, Shorea affinis, S. gardneri,

Litsea gardneri, and Palaquium rubiginosum

Subcanopy

Apodytes dimidiata, Artocarpus nobilis, Calophyllum walked, Caryota urens, Cinnamomum ovalifolium, Crypto-

carya wightiana, Dillenia triquetra, Elaeocarpus amoenus, Eugenia mabaeoides, Garcinia quaesita, Gordonia spe-

ciosa, Madhuca moonii, Mesuaferrea, Oncosperma fasciculatum, Schumacheria alnifolia, Stemonoporus gardneri,

S. oblongifolia, Syzygium firmum, and S. turbinatum

Climbers

Calamus thwaitesii, Cosinium fenestratum, Cyclea peltata, Ereycinetia walked, Rubus rugosus, and Smilax

perfoliata

Understory

Acronychia pedunculata, Agrostistachys coriacea, Alpinia abundiflora, Amomum echinocarpum, Amomum masti-

catorium, Amorphophallus paeoniifolius, Arundina graminifolia, Calanthes sp., Cinnamomum verum, Clusia rosea,

Cyathea crinita, Hedychium coronarium, Hortonia ovalifolia, Ipsea speciosa, Macaranga indica, Neolitsea cassia,

Osbeckia aspera, Osbeckia lantana, Rhodomyrtus tomentosa, Strobilanthes sp., Syzygium cordifolium, Syzygium

revolutum, and Utriculada striatula

Results and discussion

New record for D. kotagamai

We report the occurrence of the Endangered, rare, and

endemic D. kotagamai (Fernando and Day aw ansa 1994)

from Uda Maliboda forest (Uda Maliboda Trail) in the

northwest region of the Samanala Nature Reserve (SNR

= Peak Wilderness Sanctuary). According to Fernando

et al. (1994), this species is distinguished from other

Duttaphrynus species known from Sri Fanka and south-

ern India by combination of the following characters:

prominent parietal ridges on the head; long and narrow

unlobulated parotoid glands; most areas of the anterior

back are smooth; warts present on upper flank, supraor-

bital, and parietal ridges; tips of digits and tips of spinous

warts black; first Anger slightly longer than second Anger

(Fernando et al. 1994). Coloration in life is described as:

orange-brown on dorsal surface mottled with dark brown

(juveniles dorsal color is light golden); light cross band

between eyes and distinct dark cross band on forearm,

forefoot, tarsus, and tibia; less distinct cross band on up-

per arm and femur; lower jaw with alternate dark and

light markings; ventral surface whitish mottled with dark

brown, especially over sternum.

Eleven D. kotagamai were encountered during our

survey. These toads were only found in primary forest

and absent from human-disturbed areas. Except for one

specimen, all were found within ~10 m of a small stream.

(Fig. 3), and all but four individuals were observed at

night. Three individuals from Uda Maliboda measured:

two males SVF 32.6 mm, 35.2 mm, and a female SVF

38.5 mm. We also found D. kotagamai in another previ-

ously unknown locality on an adjacent mountain in De-

raniyagala in Kegalle district (Table 2). This mountain

is located about five km north of Uda Maliboda. There

are no previous records of D. kotagamai from the Uda

Maliboda Trail (SNR; see De Silva 2009; Dutta and

Manamendra-Arachchi 1996; lUCN-SF 2011; Mana-

mendra-Arachchi and Pethiyagoda 2006; Goonatilake

and Goonatilake 2001). The Uda Maliboda locality is

approximately six km (direct distance) from “Eratne”

(Kuru river basin), the nearest published location. The

direct distance between the onymotope and the new loca-

tion is about 80 km. All of these areas have closed cano-

pies with wet and cool habitats (Fig. 4).

Figure 2. View of forest in Uda Maliboda (larger water resource in the SNR).

January 2012 | Volume 5 | Number 2 | e38

amphibian-reptile-conservation.org

054

Peabotuwage et al.

Figure 3. Cascade habitat: shrub mixed with riverine forest

patch.

Figure 4. Inside forest: tall trees, mixed vegetation with good

leaf litter.

Based on the infrequent calls heard during our sur-

vey periods this species is presumably rare in Uda Mali-

boda. It is aggressive when handled and releases a low-

pitched distress call “crick, crick, crick...”. With two

new locations and a subsequent range extension, we can

trace the probable distribution of D. kotagamai prior to

fragmentation. The new locations indicate a larger distri-

bution than previously concluded. As a result of severe

fragmentation and habitat degradation in the area, local

extinctions of previous populations have likely occurred

in the past with current populations known only from a

few isolated primary forest patches.

Herpetofaunal diversity

During the study we encountered 34 amphibian species

representing 15 genera and seven families (Table 3).

Among those genera. Adenomus, Lankanectes, Nannoph-

rys, and Taruga are endemic to Sri Lanka. Our results

show that at least 31% of Sri Lanka’s extant amphib-

ians occur in the Uda Maliboda area (Fig. 5). Twenty-

six of the 34 species encountered (76%) are endemic,

five (14%) are considered Near Threatened, four (11%)

are Vulnerable, and ten (29%) are classified as Endan-

gered (lUCN-SL and MENR-SL 2007). Families with

the greatest number of endemic species include Rhaco-

phoridae (16 species) and Dicroglossidae (six species),

while the family Ichthyophiidae, Ranidae (two species

each) and Nyctibatrachidae (one species) show the low-

est rates of endemism. When considering the 34 species

by their primary mode of living, 15 (44.1%) were arbo-

real, 10 (29.4%) terrestrial, seven (20.6%) aquatic, and

two (5.9%) fossorial species.

Most amphibian species observed after brief peri-

ods of rain since many species frequently use temporary

pools created by these showers. Two large streams course

forest acting as barriers that restrict some species to par-

ticular habitats. Among the most commonly encountered

amphibians were Pseudophilautus folicola, found on

low growing woody vegetation near water bodies under

closed canopy, and Fejervarya kirtisinghei, occurred

near water bodies lacking canopy. Four Endangered and

endemic highland species: P. alto (1,890-2,135 m eleva-

tion), P. asankai (810-1,830 m), P. femoralis (1,600-

2,135 m), and Taruga eques (1,750-2,300 m; Manamen-

dra-Arachchi and Pethiyagoda 2006) were encountered

at this study site, approximately 700 m elevation (lowest

elevation ever recorded for these species).

We report a range extension for PseudophUautus

sarasinorum, an Endangered species previously known

only from the following localities: Peradeniya (07° 16’

N, 80°37’ E; Onymotope); Bogawanthalawa-Balangoda

road (near 25th km post), elevation 1,300 m (06°45’ N,

80°2’ E); Corbett’s Gap, elevation 1,000 m (07°22’ N,

80°50’ E); Hunnasgiriya, elevation 367 m (07°23’ N,

80°41’ E); Agra Arboretum, elevation 1,555 m (06°50’

January 2012 | Volume 5 | Number 2 | e38

amphibian-reptile-conservation.org

055

Uda Maliboda trail and a preliminary herpetofaunal checklist

Table 2. Description of the 1 1 observed D. kotagamai individu-

als during the study period from Uda Maliboda.

Date

Sex

Micro-habitat

18 January 2009

Male

Mid-stream boulder

Male

Forest floor with leaf litter

Female

Stream-bank boulder

17 April 2009

Female

Rock crevice

Male

Stream-bank boulder

25 December 2009

Male

Stream-bank

07 May 2010

Male

Stream-bank

Male

Stream-bank

22 August 2010

Female

Forest floor with leaf litter

Male

On footpath

03 October 2011

Male

Stream-bank boulder

N, 80°40’ E; Manamendra-Arachchi and Pethiyagoda

2005) . Sumida et al. (2007) suggested the Sri Lankan

population of F. limnocharis (in Dutta and Manamendra-

Arachchi 1996; Manamendra-Arachchi and Pethiyagoda

2006) could be F. syhadrensis. However, recent molecu-

lar evidence revealed the Sri Lankan population of F. cf.

syhadrensis is a separate and unnamed population be-

longing to a unique clade, together with F granosa and

F pierrei (Kotaki et al. 2010). Therefore, we refrain from

referring to the third Fejervarya species in Sri Lanka

as F. limnocharis (in Dutta and Manamendra-Arachchi

1996; Manamendra-Arachchi and Pethiyagoda 2006)

and instead refer it to as F. cf. syhadrensis.

Lifty-nine species of reptiles representing 37 gen-

era from 1 1 families were recorded during these surveys

(Table 4). Among those genera Aspidura, Balanophis,

Ceratophora, Cercaspis, Haplocercus, Lankascincus,

Lyriocephalus, and Nessia are considered endemic to

Sri Lanka. Twenty-eight percent of Sri Lanka’s extant

reptiles were recorded in the study area (Lig. 5) includ-

ing 28 species of lizards and 3 1 species of snakes. Of

these 59 reptile species 32 (54%) are endemic, six (10%)

Data Deficient, ten (17%) Near Threatened, five (8%)

Vulnerable, and four (7%) Endangered (lUCN-SL and

MENR-SL 2007). Lamilies with the greatest species rep-

resentation include Colubridae (17 species), Scincidae

(11 species), and Gekkonidae (nine species), while the

least represented family were Cylindrophidae, Pythoni-

dae, and Typhlopidae (one species each). The highest

number of endemic species were in the family Scincidae

(nine species) and Colubridae (seven species), while the

lowest number were in Cylindrophidae, Elapidae, and

Typhlopidae (one species each). When considering the

59 species by primary mode of living: 24 (40.7%) were

terrestrial, 21 (35.6%) arboreal, 11 (18.6%) fossorial, and

three (5.1%) aquatic species.

Among the reptiles, Otocryptis wiegmanni, Lankas-

cincus greeri, Dendrelaphis schokari, and Hypnale zara

were the most commonly encountered species in and

around footpaths. One unidentified species from the ge-

nus Cyrtodactylus was recorded during this survey and

may be new to science. Several species of lizards {Cne-

maspis scalpensis, C. silvula, Hemiphyllodactylus typus,

Eutropis beddomii, and Varanus bengalensis) and snakes

(Boiga beddomei, Cercaspis carinatus, Haplocercus cey-

lonensis, Aspidura guentheri, Balanophis ceylonensis,

and Typhlops mirus) are noteworthy records. The Uda

Maliboda forest area also supports three highly venom-

ous snakes: Bungarus ceylonicus (Sri Lanka krait), Da-

boia russelii (Russell’s viper), and Naja naja (Indian co-

bra). Hence, both venomous and non- venomous snakes

are frequently killed in this area due to fear and igno-

rance as a precautionary measure against snakebites. We

failed to record any turtle species in the area, possibly

due to low water temperatures in streams.

Figure 5. Comparison of amphibian (left) and reptile (right) diversity of Uda Maliboda area with rest of the Sri Lankan species

(Abbreviations: NOSL - total number of species in Sri Lanka; NOU - total number of species in Uda Maliboda; ENSL - number

of endemic species to Sri Lanka; ENU - number of endemic species in Uda Maliboda; TRSL - number of threatened species in Sri

Lanka and TRU - number of threatened species in Uda Maliboda).

January 2012 | Volume 5 | Number 2 | e38

amphibian-reptile-conservation.org

056

Peabotuwage et al.

Table 3. Checklist of amphibian species in the Uda Maliboda

area (Abbreviations: E - endemic; EN - Endangered; VU -

Vulnerable; NT - Near Threatened).

Family and species name

Common name

Bufonidae

Adenomus kelaartii

Kelaart’s dwarf toad ®

Duttaphrynus kotagamai

Kotagama’s dwarf toad

Duttaphrynus melanostictus

Common house toad

Microhylidae

Kaloula taprobanica

Common bull frog

Microhyla rubra

Red narrow mouth frog

Ramanella nagaoi

Nagao’s pugsnout frog

Ramanella obscura

Green-brown pugsnout frog

Nyctibatrachidae

Lankanectes corrugatus

Corrugated water frog ®

Dicroglossidae

Euphlyctis cyanophlyctis

Skipper frog

Euphlyctis hexadactylus

Sixtoe green frog

Eejervarya kirtisinghei

Montain paddy field frog ®

Eejervarya cf. syhadrensis

Common paddy field frog

Hoplobatrachus crassus

Jerdon’s bull frog

Nannophrys ceylonensis

Sri Lanka rook frog

Rhacophoridae

Pseudophilautus abundus

Labugagama shrub frog ®

Pseudophilautus alto

Horton plains shrub frog

Pseudophilautus asankai

Asanka’s shrub frog

Pseudophilautus cavirostris

Hollow snouted shrub frog

Pseudophilautus femoralis

Leafnesting shrub frog

Pseudophilautus folicola

Leaf dwelling shrub frog

Pseudophilautus hoipolloi

Anthropogenic shrub frog ®

Pseudophilautus popularis

Common shrub frog ®

Pseudophilautus reticulatus

Reticulated-thigh shrub frog

Pseudophilautus rus

Kandiyan shrub frog

Pseudophilautus sarasinorum

Muller’s shrub frog ® ™

Pseudophilautus sordidus

Grubby shrub frog

Pseudophilautus stictomerus

Orange-canthal shrub frog

Polypedates cruciger

Common hour-glass tree frog ®

Taruga eques

Mountain tree frog ™

Taruga longinasus

Long-snout tree frog ™

Ranidae

Hylarana aurantiaca

Small wood frog

Hylarana temporalis

Common wood frog

Ichthyophiidae

Ichthyophis glutinosus

Common yellow-band caecilian ®

Ichthyophis pseudangularis

Lesser yellow-band caecilian

Threats and conservation

We believe the high diversity in wet zone forest habitats

is due mainly to availability of abundant suitable micro-

habitat features (e.g., tree holes, caves, tree barks, rock

boulders, crevices, water holes, decaying logs, loose soil,

and other small niches) which create favorable environ-

mental conditions for herpetofauna. According to our re-

sults, Uda Maliboda area has a rich herpetofaunal diver-

sity and endemism compared with other wet zone forests

in Sri Lanka. A large number of people including tourists,

devotees, and laborers annually visit Adams Peak via

Uda Maliboda Trail located within the SNR. As a result

endemic and Threatened species, like many other fauna,

are seriously affected by increasing pressure caused by

habitat loss and degradation in montane forests, lower

montane forests, and marshes. Major threats identified in-

clude illegal timber harvesting, illegal human encroach-

ment, slash and bum forest clearing for human settlement

and monoculture plantations (especially for tea cultiva-

tion), and gem mining. According to interviews with il-

legal timber harvesters, some rare tree species may be

new to science are being harvested. Therefore, a further

comprehensive study of fiora is recommended.

Present human activities, the most severe being the

constmction of a hydroelectric power plant, continue to

degrade and erode the remaining vestiges of this lush pri-

mary forest. Additionally, garbage (polythene) disposal

along the Uda Maliboda Trail by visitors and devotees is

a threat that must be duly monitored by the Department

of Wildlife Conservation (DWC) and the Forest Depart-

ment (FD) of Sri Lanka. The Young Zoologists’ Associa-

tion (YZA) together with the Central Environmental Au-

thority (CEA) has conducted annual polythene removal

programs on other trail (Hatton) of SNR for the past 10

years. This has prompted other Government institutions

and non-governmental organizations to engage in similar

activities. We recommend that such programs be initiated

on this trail in order to prevent further degradation of this

lush forest.

Some human-altered landscapes such as tea planta-

tions and Pinus, Eucalyptus, Cyprus, and Casuarina for-

est plantations are located in the foothills of the SNR.

Most of these altered landscapes can be found up to

about 800 m in elevation. There is an ongoing hydroelec-

tric power plant development project in the study area

(Fig. 6) and increased road traffic further threatens the

area’s fauna. Since a considerable area of the forest is

altered by human activity, herpetofauna face increased

threats because, in general, they are often highly sensi-

tive to even slight environmental changes (e.g., McCal-

lum 2007; Pough et al. 2004; Spellerberg 1991). Thus,

the identification and designation of forest reserves on

the perimeter of the SNR could function as suitable buf-

fer zones. Additionally, public awareness programs are

needed to help guide local people and policy makers de-

January 2012 I Volume 5 | Number 2 | e38

amphibian-reptile-conservation.org

057

Uda Maliboda trail and a preliminary herpetofaunal checklist

Table 4. Checklist of reptile species in Uda Maliboda area (Abbreviations: E - endemic; EN - Endangered; VU - Vulnerable; NT

- Near Threatened; DD - Data Deficient.

Family and

species name

Common name

Agamidae

Calotes calotes

Green garden lizard

Calotes liolepis

Whistling lizard

Calotes versicolor

Common garden lizard

Ceratophora aspera

Rough horn lizard ® ™

Lyriocephalus scutatus

Lyre-head lizard ^

Otocryptis wiegmanni

Sri Lankan kangaroo lizard

Gekkonidae

Cnemaspis scalpensis

Gannoruva day geeko °°

Cnemaspis silvula

Forest day geeko ®

Cyrtodactylus cf. subsolanus

Forest geeko sp.

Geckoella triedrus

Spotted bowfinger gecko

Gehyra mutilata

Four-elaw geeko

Hemiphyllodactylus typus

Slender geeko

Hemidactylus depressus

Kandyan geeko ®

Hemidactylus frenatus

Common house geeko

Hemidactylus parvimaculatus

Spotted house geeko

Scincidae

Eutropis beddomii

Beddome’s stripe skink®’®'^

Eutropis carinata

Common skink

Eutropis macularia

Bronzegreen little skink

Eutropis madaraszi

Spotted skink

Lankascincus dorsicatenatus

Catenated lankaskink ®

Lankascincus fallax

Common lankaskink®

Lankascincus gansi

Gans’s lankaskink ®’

Lankascincus greeri

Greer’s lankaskink ®

Lankascincus munindradasai

Munidradasa’s lankaskink ®’ °°

Lankascincus sripadensis

Peakwildemess lankaskink ®’ °°

Nessia burtonii

Three toed snakeskink ®

Varanidae

Varanus bengalensis

Land monitor

Varanus salvator

Water monitor

Pythonidae

Python molurus

Indian python

Cylindrophidae

Cylindrophis maculatus

Sri Lanka pipe snake ®’

Colubridae

Ahaetulla nasuta

Green vine snake

Ahaetulla pulverulenta

Brown vine snake

Boiga barnesii

Barnes’s eat snake

Boiga beddomei

Beddoms eat snake °°

Boiga ceylonensis

Sri Lanka eat snake

Cercaspis carinatus

Sri Lanka wolf snake ®’

Coeloganthus helena

Trinket snake

Dendrelaphis bifrenalis

Boulenger’s bronze baek ®

Dendrelaphis caudolineolatus

Gunther’s bronze baek

Family and

species name

Common name

Colubridae (cont.)

Dendrelaphis schokari

Common bronze baek ®

Haplocercus ceylonensis

Blaek spine snake

Lycodon aulicus

Common wolf snake

Lycodon striatus

Shaw’s wolf snake

Oligodon calamarius

Templeton’s kukri snake ®

Oligodon sublineatus

Dumerul’s kuki snake ®

Ptyas mucosa

Rat snake

Sibynophis subpunctatus

Jerdon’s polyodent

Natricidae

Amphiesma stolatum

Buff striped keelbaek

Aspidura guentheri

Ferguson’s roughside ®’

Balanophis ceylonensis

Sri Lanka keelbaek ®’

Atretium schistosum

Olive keelbaek

Xenochrophis asperrimus

Cheekered keelbaek ®

Typhlopidae

Typhlops mirus

Jan’s blind snake ®

Elapidae

Bungarus ceylonicus

Sri Lanka krait ®’

Naja naja

Indian eobra

Viperidae

Daboia russelii

Russell’s viper

Hypnale hypnale

Merrem’s hump nose viper

Hypnale zara

Zara’s hump-nosed viper ®

Trimeresurus trigonocephalus

Green pit viper ®

velop agendas that consider the importance of herpeto-

fauna in maintaining a balanced and healthy ecosystem.

There is no doubt that SNR provides habitat for a

high number of amphibian and reptiles species (many

endemic and Threatened). We affirm that it is one of

the most important herpetofaunal diversity areas in Sri

Lanka, especially when considering the future conserva-

tion of endemic and threatened herpetofauna. Sri Lanka

is known as an important herpetofaunal global hotspot

(Bossuyt et al. 2004; Gunawardene et al. 2007; Meegas-

kumbura et al. 2002; Pethiyagoda 2005) and harbors an

unusually high number of endemic species. Therefore,

scientists and policy makers are strongly encouraged to

make efforts conducting further research on other fau-

nal groups, vegetation, and the forest’s ecosystem as a

whole. Furthermore, preserving the valuable herpetofau-

nal resources of the Uda Maliboda Trail is paramount to

the conservation of global biological diversity.

Acknowledgments. — We would like to express our

sincere gratitude to Thasun Amarasinghe (Taprobanica)

for reviewing the earlier draft of the manuscript. We

also thank Mendis Wickramasinghe (HFS), Aruna Ka-

January 2012 | Volume 5 | Number 2 | e38

amphibian-reptile-conservation.org

058

Peabotuwage et al.

Figure 6. Hydroelectric power plant (note: concrete wall built

across the steam and concrete particles dump into the steam).

runathilake, Nadeesh Gamage, Mahesh De Silva (YZA),

Prof. Deepthi Yakandawala, Dr. Suranjan Fernando (Uni-

versity of Peradeniya), and other members of the Young

Zoologists’ Association of Sri Lanka (YZA) for various

help with this study. Villagers in the Uda Maliboda area

are acknowledged for their cooperation, sharing their ob-

servations, and logistic support. Finally, we would like

to give our special thanks to John Rudge, Daniel Fogell,

Kanishka Ukuwela, and Craig Hassapakis (ARC) for

reviewing the initial daft of the manuscript and making

improvements.

Literature cited

Ashton MS, Gunatileke CVS, De ZoysaN, Dassanayake MD, Gu-

NATiLEKE lAUN, WiJESUNDARA S. 1997. A Field Guide to the Com-

mon Trees and Shrubs of Sri Lanka. Wildlife Heritage Trust of Sri

Lanka, Colombo. 432 p.

Bambaradeniya CNB, Perera MSI, Perera WPN, Wickramasinghe

LJM, Kekulandala LDCB, Samarawickrema VAP, Fernando

RHSS, Samarawickrema VAMPK. 2003. Composition of faunal

species in the Sinharaja world heritage site in Sri Lanka. Sri Lan-

ka Forester 26(3):21-40.

Bossuyt F, Meegaskumbura M, Beenaerts N, Gower DJ, Pethiya-

GODA R, Roelants K, Mannaert a, Wilkinson M, Bahir MM,

Manamendra-arachchiK,NgPKL, Schneider CJ, OommenOV,

Milinkovitch MC. 2004. Local endemism within the Western

Ghats - Sri Lanka Biodiversity Hotspot. Science 306:479-481.

Bossuyt F, Meegaskumbura M, Baenerts N, Gower DJ, Pethiya-

GODA R, Roelants K, Mannaert A, Wilkinson M, Bahir MM,

Manamendra-Arachchi K, Ng PKL, Schneider CJ, Oomen OV,

Milinkovitch MC. 2005. Biodiversity in Sri Lanka and Western

Ghats - Response. Science 308:199.

Crump ML, Scott NJ. 1994. Visual encounter surveys. In: Measur-

ing and Monitoring Biological Diversity: Standard Methods for

Amphibians. Editors, Heyer RW, Donnelly MA, McDiarmid RW,

Hayek LC, Foster MS. Smithsonian Institution Press, Washing-

ton, D.C. 84-92.

De Silva A. 2009. Amphibians of Sri Lanka: A Photographic Guide to

Common Frogs, Toads and Caecilians. Published by the author.

82 plates -i- 168 p.

DuttaSK, Manamendra-Arachchi KN. 1996. The Amphibian Fauna

of Sri Lanka. Wildlife Heritage Trust of Sri Lanka, Colombo, Sri

Lanka. 230 p.

Fernando P, Dayawansa, N, Siriwardene M. 1994. Bufo kotagamai,

a new toad (Bufonidae) from Sri Lanka. Journal of South Asian

Natural History 1(1):119-124.

Frost DR, Grant T, Faivovich J, Bain RH, Haas A, Haddad CFB, de

S aR, ChanningA, WilkinsonM, Donnellan S C, RaxworthyC J,

Campbell JA, Blot-to BL, Moler P, Drewes RC, Nussbaum RA,

Lynch JD, Green DM, Wheeler WC. 2006. The amphibian tree

of life. Bulletin of American Museum of Natural History 297:1-

370.

Goonatilake WLDPTS de A, Goonatilake MRMPN. 2001. New

sight records of Bufo kotagamai (Anura: Bufonidae) from Adavi-

kanda in Eratne and Delwala proposed forest Reserve. Sri Lanka

Naturalist 4(4):55-56.

GunatillekeIAUN, Gunatilleke CVS. 1 990. Distribution offloristic

richness and its conservation in Sri Eanka. Conservation Biology

4(1):21-31.

GunawardeneNR, Daniels AED, Gunatilleke IAUN, Gunatilleke

CVS, Karunakaran PV, Nayak KG, Prasad S, Puyravaud P, Ra-

MESH BR, SuBRAMANiAN KA, Vasanthy G. 2007. A brief overview

of the Western Ghats-Sri Eanka biodiversity hotspot. Current Sci-

ence 93(11):1567-1572.

lUCN-SE 2011. Threatened Biodiversity (A coffee table book). lUCN

Sri Lanka country office, Colombo, Sri Lanka. 212 p.

lUCN-SL, MENR-SL. 2007. The 2007 Red List of Threatened Fauna

and Flora of Sri Lanka. lUCN Sri Eanka, Colombo, Sri Eanka.

148 p.

KarunarathnaDMSS, AmarasingheAAT, BandaraIN. 201 1 . Asur-

vey of the avifaunal diversity of Samanala Nature Reserve, Sri

Eanka, by the Young Zoologists’ Association of Sri Eanka. Bind-

ing Asia 15:84-91.

Kotaki M, Kurabayashi a, Matsui M, Kuramoto M, Djong TH,

SuMiDA M. 2010. Molecular phylogeny for the diversified frogs

of genus Fejervarya (Anura: Discoglossidae). Zoological Science

27(5):386-395.

McCallum me. 2007. Amphibian decline or extinction? Current de-

clines dwarf background extinction rate. Journal of Herpetology

41(3):483-491.

Maduwage K, Silva a, Manamendra-Arachchi K, Pethiyagoda R.

2009. A taxonomic revision of the South Asian hump-nosed pit

vipers (Squamata: Viperidae: Hypnale). Zootaxa 2232:1-28.

Manamendra-Arachchi K, Pethiyagoda R. 2005. The Sri Eankan

shrub-frogs of the genus Philautus Gistel, 1848 (Ranidae: Rha-

cophorinae) with description of 27 new species. In: Contribution

to Biodiversity Exploration and Research in Sri Lanka. Editors,

Yeo DCJ, Ng PKL, Pehiyagoda R. The Raffles Bulletin of Zool-

ogy, Supplement 12:163-303.

Manamendra-Arachchi K, Pethiyagoda R. 2006. SriLankawe Ub-

hayajeeween [Amphibians of Sri Lanka]. Wildlife Heritage Trust

of Sri Lanka, Colombo, Sri Lanka. 88 plates -i- 440 p. (Text in

Sinhala).

January 2012 | Volume 5 | Number 2 | e38

amphibian-reptile-conservation.org

059

Uda Maliboda trail and a preliminary herpetofaunal checklist

Figure 7. Duttaphrynus kotagamai (Male; Endangered).

Figure 8. Lankanectes corrugatus (relict).

Figure 9. Psedophilautus femoralis (Endangered).

Figure 10. Psedophilautus reticulates (Endangered).

Figure 11. Pseudophilautus alto (Endangered).

Figure 12. Pseudophilautus sarasinorum (Endangered).

Figure 13. Ramanella nagaoi (Vulnerable).

Figure 14. Taruga longinasus (Endangered).

January 2012 | Volume 5 | Number 2 | e38

amphibian-reptile-conservation.org

060

Peabotuwage et al.

Figure 15. Oligodon calamarius (Vulnerable).

Figure 16. Dendrelaphis schokari (Endemic).

Figure 17. Amphiesma stolatum (red variety).

Figure 18. Trimeresurus trigonocephalus (plain variety).

Figure 19. Hemidactylus depressus (endemic).

Figure 20. Unidentified Cyrtodactylus cf. subsolanus.

Figure 21. Lankascincus greeri (endemic).

Figure 22. Eutropis macularia (common).

January 2012 | Volume 5 | Number 2 | e38

amphibian-reptile-conservation.org

061

Uda Maliboda trail and a preliminary herpetofaunal checklist

Figure 23. Ceratophora aspera (Endangered).

Man AMENDRA- Arachchi K, B atuwita S , PethiyagodaR. 2007 .Atax-

onomical revision of the Sri Lankan day-geckos (Reptilia: Gek-

konidae: Cnemaspis), with description of new species from Sri

Lanka and India. Zeylanica 7(1):9-122.

Meegaskumbura M, Bossuyt F, Pethiyagoda R, Manamendra-

Arachchi K, Bahir M, Milinkovitch MC, Schneider CJ. 2002.

Sri Lanka: An amphibian hotspot. Science 298(5592):379.

Meegaskumbura M, Meegaskumbura S, Bo watte G, Manamendra-

Arachchi K, Pethiyagoda R, Hanken J, Schneider CJ. 2010.

Taruga (Anura: Rhacophoridae), A new genus of Form-nesting

tree frogs endemic to Sri Lanka. Ceylon Journal of Science (Bio

science) 39(2):75-94.

Mittermeier RA, Gil PR, Hoffman M, Pilgrim J, Brooks T, Mitter-

MEiER CG, Lamoreux J, DA FoNSECA GAB. 2004. Hotspots Revis-

ited: Earth ’5 Biologically Richest and Most Threatened Terrestrial

Ecoregions. CEMEX, Mexico City and Conservation Internation-

al, Washington, D.C. 164 p.

Myers N, Mittermeier RA, Mittermeier CG, Fonseca GAB, Kent

J. 2000. Biodiversiy hotspots for conservation priorities. Nature

403:853-858.

Pethiyagoda R. 2005. Exploring Sri Eanka’s biodiversity. In: Con-

tribution to Biodiversity Exploration and Research in Sri Lanka.

Editors, Yeo DCJ, Ng PKL, Pehiyagoda R. The Raffles Bulletin of

Zoology, Supplement 12:1-4.

PouGH FH, Andrews RM, C adle JE, Crump ML, S avitzkyAH, W ells

KD. 2004. Herpetology. 3rd Edition. Pearson Prentice Hall, San

Erancisco, USA. 726 p.

Senanayake FR, Soule M, Senner JW. 1977. Habitat values and en-

demicity in the vanishing rainforest of Sri Eanka. Nature 265:35 1-

354.

Senaratna EK. 2001. A Check List of the Elowering Plants of Sri

Lanka. National Science Eoundation of Sri Eanka, Colombo. 342

P-

SoMAWEERA R. 2006. Sri Lankawe Sarpayan [Snakes of Sri Eanka].

Wildlife Heritage Trust of Sri Eanka, Colombo Sri Eanka. 88

plates -I- 440 p. (Text in Sinhala).

SoMAWEERA R, SoMAWEERA N. 2009. Lizards of Sri Lanka: A Colour

Guide with Eield Keys. Edition Chimaira, Erankfurt am Main,

Germany. 303 p.

Figure 24. Calotes liolepis (Vulnerable).

Spellerberg IE. 1991. Monitoring Ecological Changes. Cambridge

University Press, Cambridge, UK. 334 p.

SuMiDA M, Kotaki M, Islam MM, Djong TH, Igawa T, Kondo Y,

Matsui M, De Silva A, Khonsue W, Nishioka M. 2007. Evolu-

tionary relationships and reproductive isolating mechanisms in

the Rice frog (Eejervarya limnocharis) species complex from Sri

Eanka, Thailand, Taiwan and Japan, inferred from mtDNA gene

sequences, allozymes, and crossing experiments. Zoological Sci-

ence 24:547-562.

UNITEDNAriONSEDUCAriONAL,SciENTIFICANDCULTURALORGANIZAriON

(UNESCO). 2011. UNESCO Headquarters, 7, Place de Eontenoy,

75352, Paris, 07 SP, Prance. [Online]. Available: http://whc.unes-

co.org/en/list/1203/documents/ [Accessed: 25 November 2011].

Vogel G, Rooijen JV. 201 1 . A new species of Dendrelaphis (Serpen-

tes: Colubridae) from the Western Ghats - India. Taprobanica

3(2):77-86.

WiCKRAMASiNGHE LJM, Rodrigo R, Dayawansa N, Jayantha UED.

2007a. Two new species of Lankascincus (Squamata: Scincidae)

from Sripada. Zootaxa 1612:1-24.

WiCKRAMASiNGHE LJM, MuNiNDRADASA DAI. 2007b. Rcvicw of the

genus Cnemaspis Strauch, 1887 (Sauria: Gekkonidae) in Sri Lan-

ka, with the description of five new species. Zootaxa 1490:1-63.

WiJESiNGHE MR, Dayawansa PN. 2002. The amphibian fauna at two

altitudes in the Sinharaja rainforest, Sri Lanka. Herpetological

Journal 12:175-178.

Manuscript received: 30 November 2011

Accepted: 26 December 2011

Published: 18 January 2012

January 2012 | Volume 5 | Number 2 | e38

amphibian-reptile-conservation.org

062

Peabotuwage et al.

INDIKAPEABOTUWAGE is a botanist working at the Depart-

ment of Botany, University of Peradeniya and has great skill in

botanical illustrating. He is a member of the Young Zoologists

Association (YZA) and president of the research committee.

During his career, he has participated in several national and

international training programs. At present, he works on sev-

eral plant based research projects and conserving the vanishing

biodiversity in Sri Lanka.

DINAL SAMARASINGHE is a Sri Lankan herpetologist,

wildlife photographer, and member of the Young Zoologists’

Association (YZA) based at the National Zoological Gardens

of Sri Lanka. His research is mainly focused on territoriality,

aggressive behavior, and vocal communication in anurans.

Presently, he leads a study on systematics, distribution patterns,

and ecology of the genus Varanus in India and Sri Lanka. Dinal

also works as a venom extractor at the Snake Venom Research

Laboratory and Herpetarium (SVRLH), Laculty of Medicine,

University of Colombo.

NUWAN BANDARA is a graduate from the University of Per-

adeniya, and his scientific exploration of biodiversity began

with the Youth Exploration Society of Sri Lanka (YES) in late

1990. As a member and former president of YES, he is conduct-

ing biodiversity conservation and education programs for the

Sri Lankan community. His specific fields of research interest

are ecosystem services, community-based conservation, tradi-

tional agricultural practices, ethnobotany, and local biodiversity

and behavioral ecology of herpetofauna and other wild fauna.

NIRMALA PERERA is a naturalist and has had a special in-

terest in amphibians and reptiles ever since his childhood. He

conducts various conservation events on biodiversity restora-

tion and education programs for the local community and as

an environmentalist, he is engaged in numerous snake rescue

programs. He is a member of the Young Zoologists’ Associa-

tion (YZA), National Zoological Gardens of Sri Lanka and cur-

rently works as a project manager (Human-Elephant Confiict

Program, Udawalawe) for the Born Eree Eoundation, Sri Lanka

country office.

January 2012 | Volume 5 | Number 2 | e38

amphibian-reptile-conservation.org

063

Uda Maliboda trail and a preliminary herpetofaunal checklist

MAJINTHA MADAWALA is a naturalist and conducts several

habitat restoration programs in many forest areas. He began his

career and wildlife interests in 1995 as a member of the Young

Zoologists’ Association (YZA), National Zoological Gardens

of Sri Lanka. He holds a Diploma in biodiversity management

from the University of Colombo. As a conservationist, he is

engaged in numerous snake rescue programs and funding for

ongoing research projects.

DUSHANTHA KANDAMBI is a researcher conducting and

supporting investigations on amphibians and reptiles. He is

also engaged in a captive breeding program on threatened spe-

cies and rescue events. Additionally, he promotes conservation

awareness of the importance of snake fauna among the Sri

Lankan community. He is a wildlife artist and photographer

enjoying nature.

CHAMARA AMARASINGHE is a researcher interested in

fauna and flora of Sri Lanka. He has a keen interest in freshwa-

ter ichthyofauna, butterflies, birds, marine mammals, and bats.

He is a wildlife artist and photographer engaged with the Youth

Exploration Society of Sri Lanka (YES). He started his passion

to explore much of the islands rare and endangered animals at

a very young age. Currently, he is working as a naturalist at

Jetwing Blue, a prestigious tourist hotel in Sri Lanka.

SURANJAN KARUNARATHNA is a field biologist conduc-

ing research on amphibian and reptile ecology, and promot-

ing conservation awareness of the importance of biodiversity

among the Sri Lankan community. He began his career and

wildlife research in 2000, as a member of the Young Zoologists’

Association (YZA), National Zoological Gardens of Sri Lanka.

He worked as an ecologist for the lUCN Sri Lanka county of-

fice and is an active member of many specialist groups in the

lUCN/SSC.

January 2012 | Volume 5 | Number 2 | e38

amphibian-reptile-conservation.org

064

Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any

medium, provided the original author and source are credited.

Amphibian & Reptiie Conservation 5(2):65-80.

Herpetofaunal diversity and distribution in Kalugaia proposed

forest reserve, Western province of Sri Lanka

' 3W. MADHAVA S. BOTEJUE AND ^JAYANTHA WATTAVIDANAGE

^Taprobanica Nature Conservation Society, 150/6, Stanley Thilakaratne Mawatha, Nugegoda, SRI LANKA ^Department of Zoology, Faculty of

Natural Sciences, The Open University of Sri Lanka, SRI LANKA

Abstract . — Kalugaia Proposed Forest Reserve (KPFR) is a primary lowland tropical rain forest, sur-

rounded by secondary forest and vegetation disturbed by human activities such as cultivation,

logging, and the collection of firewood. Herpetofaunal communities of selected different habitats

(closed forest, forest edge, home gardens, and cultivations) were assessed and distribution pat-

terns were compared. A total of 24 amphibian species (63% endemic and 33% Threatened) and 53

reptile species (38% endemic and 30% Threatened) were recorded. Overall, 763 individual amphib-

ians and 1032 individual reptiles were recorded in this forest area. Reptilian distribution patterns

are similar to amphibian distribution patterns, with the highest diversity in the closed forest and the

lowest diversity in cultivations. We did not observe an effect of forest edge (edge effect) in amphib-

ian and reptile diversity, except for forest edge and cultivations for reptiles. Adverse human activi-

ties such as improper agriculture practices, logging, and waste disposal have led to deforestation

and habitat loss in KPFR.

Key words. Amphibians, reptiles, conservation, ecology, habitats, rain forest, Sri Lanka, threats

Citation: Botejue WMS, Wattavidanage J. 2012. Herpetofaunal diversity and distribution in Kalugaia proposed forest reserve, Western province of Sri

Lanka. Amphibian & Reptile Conservation 5(2):65-80 (e39).

Introduction

Recent research has demonstrated the uniqueness of Sri

Lankan fauna and its distinctness from the Indian main-

land (Bossuyt et al. 2004, 2005; Helgen and Groves

2005). This is particularly true of the herpetofaunal

assemblage (Bossuyt et al. 2004; Meegaskumbura et

al. 2002). There are 110 species of amphibians in Sri

Lanka, which belong to seven families and 19 genera

with 95 (86%) endemic species. (Fernando et al. 2007;

Frost 2008; Manamendra-Arachchi and Pethiyagoda

2006; Meegaskumbura et al. 2007; Meegaskumbura et

al. 2009; Meegaskumbura et al. 2010; Meegaskumbura

and Manamendra-Arachchi 2011). The reptile fauna con-

sists of 210 species, including 120 (57%) endemic spe-

cies, representing 24 families and 82 genera. (Bauer et

al. 2007; Batuwita and Pethiyagoda 2007; de Silva 2006;

Gower and Maduwage 2011; Maduwage et al. 2009; Ma-

namendra-Arachchi et al. 2006; Manamendra-Arachchi

et al. 2007; Smith et al. 2008; Somaweera 2006; Wick-

ramasinghe and Munindradasa 2007 ; Wickramasinghe et

al. 2009).

In the present period of mass extinction of biodiver-

sity (Achard et al. 2002; Jenkins 2003) many species of

animals, plants, and other organisms are disappearing at

an alarming rate, primarily due to human activities such

Correspondence. Email: ^madhavabotejue® gmail.com

as deforestation (Bambaradeniya et al. 2003; Brook et al.

2003; Pethiyagoda 2005, 2007a), fire (Batuwita and Ba-

hir 2005), erosion (Hewawasam et al. 2003), agrochemi-

cal use (Pethiyagoda 1994), and lack of systematic or sci-

entific understanding (Bahir 2009; Pethiyagoda 2007b).

Although the natural forest area of Sri Lanka still consti-

tutes over 12% of the total land area (Tan 2005), human

population density of the biologically rich wet zone is

among the highest on earth (Cincotta et al. 2000). Fur-

thermore, the population growth rate is increasing around

protected areas (Wittemyer et al. 2008). Natural forests

and the biodiversity have been rapidly diminishing over

the past 100 years. The result has been the extinction of

21 species of amphibians, with 19 of these species being

from the genus Pseudophilautus (Manamendra-Arach-

chi and Pethiyagoda 2005; Meegaskumbura and Man-

amendra-Arachchi 2005; Meegaskumbura et al. 2007).

In addition, of the remaining species, 57 reptiles and 56

amphibians are considered Threatened (lUCNSL and

MENRSL 2007).

Kalugaia Proposed Forest Reserve (KPFR) is one of

the remaining few wet zone forest patches in Sri Lan-

ka and is threatened by human activities. We report the

results of a study conducted in KPFR to assess species

richness, abundance, and diversity of the herpetofauna

and to evaluate the distribution patterns among different

habitats.

January 2012 | Volume 5 | Number 2 | e39

amphibian-reptile-conservation.org

065

Botejue and Wattavidanage

Study area and habitats

The KPFR belongs to Agalawatta and Walallawita Divi-

sional Secretariat of Kaluthara District, Sri Lanka, which

lies between 6°25’-6°30’ N and 80°12’-80°16’ E (Fig.

1). The floristic structure and composition suggest KPFR

retain a considerable amount of primary forest. However

the boundaries of this forest are disturbed due to cultiva-

tion, logging, firewood collection, and consist of second-

ary and disturbed vegetation. We identified four types of

habitats as study sites: closed forest (Fig. 2), forest edge

(Fig. 3), home gardens (Fig. 4), and cultivations (Fig. 5a,

b, c).

Originally, the KPFR was an area of approxiatemly

4,630 ha when first declared a Proposed Forest Reserve

in 1992. However, due to continuous deforestation, log-

ging, agriculture practices, and illegal encroachments,

the land area has drastically reduced to about 2,907 ha

(Ranasinghe 1995). Several decades ago, KPFR was part

of the western-most extension of Sinharaja rainforest,

however, today it has been diminished to an isolated for-

est patch due to extensive deforestation and other human

activities (Kekulandala 2002; Ranasinghe 1995). The

elevation of the area ranges from 30-300 m and the ma-

jority of its precipitation originates from the southwest

monsoon (April to September) with a mean annual rain-

fall of 4000-5000 mm. The KPFR is a catchment area

for both Benthara and Kalu rivers. Average monthly tem-

perature in the region is -27.3 °C (Kekulandala 2002;

Ranasinghe 1995).

Closed forest is found deep in KPFR and on hill-

tops (Fig. 6). The major vegetation formation of this

habitat type can be classified as Doona-Dipterocarpus-

Mesua series (Ranasinghe 1995). A certain degree of

stratification can be identified in the forest, and although

an emergent layer cannot be clearly identified, at some

places the forest rises up to about 50-60 m in height and

is primarily composed of Dipterocarpus sp., Shorea sp.,

and Doona sp. The canopy layer is composed of Aniso-

phyllea cinnamomoides, Mesua sp., Valeria copallifera,

and Mangifera zeylanica, that rise to about 30-40 m. The

subcanopy is about 15-30 m high with the primary trees

being Semecarpus sp., Garcinia sp. Calophyllum sp., and

Horsfieldia iryaghedhi. The composition of the under-

story is variable, but primarily this layer is comprised

of Humboldtia laurifolia, Strobilanthes sp., Cyathea

sp., saplings of Calamus sp., and Glochidion sp. The

ground layer is mainly composed of species in the fam-

ily Poaceae and Asteraceae, as well as ground orchids.

This forest harbor a rich assemblage of climbing plants

(e.g.. Pathos sp., Entada pusaetha, and Calamus sp.) and

epiphytes. Exotic species like Alstonia macrophylla are

also found in the forest and the ground is covered with

a thick and moist decomposing leaf matter layer. A con-

siderable number of streams are located in the study area

(Fig. 7). Some areas of the forest are disturbed by well-

maintained trails (Fig. 8) and, in some places, the forest

is directly connected to cultivations.

The forest edge is the marginal area between closed

forest and home gardens or cultivations. This is highly

disturbed by human activities such as logging and fire-

wood collecting. The vegetation of this area consists of

a mixture of forest vegetation and home garden vegeta-

tion, trees such as Mesua sp., Dipterocarpus sp., Shorea

sp., Doona sp., Mangifera zeylanica, Mangifera indica,

Caryota urens, Areca catechu, Artocarpus nobilis, Ar-

tocarpus heterophyllus, Trema orientalis, Syzygium sp.,

Garcinia sp., Murraya paniculata, Elaeocarpus sp.,

Macaranga sp., Mallotus sp.; shrubs such as Ochland-

ra stridula, Osbeckia sp., Melastoma malabathricum;

climbers such as Calamus sp., and tree ferns {Cyathea

sp.). The under growth is very dense in most parts of the

forest edge, where Dicranopteris sp. and many other fern

species dominate. Species of the family Poaceae and As-

teraceae were also found in the ground layer and exotic

Figure 1. Geographical location and map of KPFR.

January 2012 | Volume 5 | Number 2 | e39

amphibian-reptile-conservation.org

066

Herpetofauna of Kalugala proposed forest reserve

Figure 2. Closed forest.

Figure 4. Home gardens.

Figure 5b. Cultivation (tea).

species like Alstonia macwphylla, Dillenia sujfruticosa.

Eucalyptus sp., Acacia sp., and Pinus sp. were present in

this habitat type.

Home garden vegetation consists of crop, shade, and

ornamental plants such as Musa sp., Mangifera indica,

Caryota urens, Areca catechu. Cocos nucifera, Carica

papaya, Artocarpus heterophyllus, Artocarpus incisus,

Syzygium sp., Garcinia sp., Elaeocarpus serratus, Ma-

caranga peltata, Manihot esculenta, Albizia sp.. Cassia

Figure 3. Forest edge.

Figure 5a. Cultivation (paddy).

Figure 5c. Cultivation (rubber).

sp., Nephelium lappaceum, Cinnamomum verum. Plume-

ria sp., Spondias sp.. Piper betle, and P. nigrum. Shrubs

consist of Melastoma malabathricum, Osbeckia octan-

dra, and exotic Lantana camara. Most home gardens are

directly associated with cultivations (Fig. 9), and thus

many herbaceous crop plants of the family Fabaceae,

Cucurbitaceae, Poaceae, and Asteraceae, and other or-

namental plants are present, as are exotic trees such as

Alstonia macrophylla and Acacia sp.

January 2012 | Volume 5 | Number 2 | e39

amphibian-reptile-conservation.org

067

Botejue and Wattavidanage

Figure 6. Forest on hilltops. Figure 9. Home gardens associated with cultivation.

The KPFR area include three main types of cultiva-

tion: paddy, tea, and rubber. Mud pools and small rivu-

lets in paddy-cultivated land provide many microhabitats

for amphibians. Around paddy and tea cultivation other

crops like banana {Musa sp.) and coconut {Cocos nu-

cifera) can be seen. Most rubber cultivations are not well

maintained and the undergrowth is high and comprised

of Dicranopteris sp., and herbaceous plants of the fam-

ily Fabaceae and Poaceae. In some locations two culti-

vations are in close proximity with one another, such as

tea and rubber, or tea and paddy (Fig. 10a, b), and in a

few locations all three cultivations can be found in close

proximity.

Materials and methods

Data collection

Dates of field study were determined using a random

number table. A total of 12 field visits were conducted for

a total of 480 hours. Visual encounter surveys and line

transects (200 m) were used for data collection, including

night visits with the aid of head lamps. Belt transects (4

X 50 m) used for data collection and observations con-

ducted 20 cm deep into the leaf litter. Quadrat sampling

(5 x5 m) was employed for habitat-specific sampling,

with quadrats being placed in pairs in every location of

each habitat type. All quadrats were surveyed once dur-

ing the day and once at night by 4-5 people moving slow-

ly inward from the periphery. Randomly placed pitfall

traps were used to sample small terrestrial reptiles where

others were hand captured. Temperature and humidity

were measured using a digital thermometer and a digital

humidity meter, respectively. Weather, cloud cover, and

canopy cover were assessed visually. In total, 24 quad-

rats, 12 line transects, and four belt transects were used,

equating a total sampling area of 1400 m^ -i- 2000 m with

equal observation time being allocated to each habitat.

Figure 7. Streams inside the forest.

Figure 8. Well maintained trails inside the forest.

January 2012 | Volume 5 | Number 2 | e39

amphibian-reptile-conservation.org

068

Herpetofauna of Kalugala proposed forest reserve

Data analysis

The Shannon- Wiener Index {H’ = -X (p- In p)] was

used to determine the diversity of species heterogene-

ity (where, H’ = species diversity, and p. = proportional

frequency of the i* species). The non-parametric Mann-

Whitney U-tQSt at the 10% significant level was used to

test differences in independent samples of amphibian and

reptile distribution among habitats.

Species Identification

All amphibian and reptile species were identified and

classified using Dutta and Manamendra-Arachci (1996),

de Silva (2009), Howlader (2011), Manamendra-Arach-

chi and Pethiyagoda (2006), Meegaskumbura et al.

(2009), Meegaskumbura et al. (2010), and Meegaskum-

bura and Manamendra-Arachchi (2011) for amphibians;

Bahir and Silva (2005), Bauer et al. (2010a and 2010b),

Das and de Silva (2005), Deraniyagala (1953 and 1955),

de Silva (1990 and 2006), Gunther (1864), Manamendra-

Arachchi et al. (2007), Pethiyagoda and Manamendra-

Arachchi (1998), Smith (1935), Somaweera (2006),

Somaweera and Somaweera (2009), Taylor (1953), and

Whitaker and Captain (2004) for reptiles. Plant species

were identified using Ashton et al. (1997), Dassanayake

and Fosberg (1980-1991), Dassanayake et al. (1994-

1995), Dassanayake and Clayton (1996-2000), Guna-

tilleke and Gunatilleke (1990), and Senaratna (2001).

The lists of Threatened species were based on the most

recent national Red List (lUCNSL and MENRSL 2007).

Results

Species richness

A total of 24 species of amphibians (representing 15

genera in 7 families) were recorded, with 15 species

(63%) being endemic, and eight (33%) being Threatened

(Table 1). A total of 53 species of reptiles (representing

38 genera and 12 families) were recorded, with 20 spe-

cies (38%) being endemic and 16 (30%) being Threat-

ened (Table 2). The greatest species richness for both

amphibians and reptiles was in closed forest, with all 24

species of amphibians being recorded there, and 45 spe-

cies (85%) of reptiles. For amphibians, 23 species (96%;

excluding Pseudophilautus reticulatus) were recorded in

forest edge, followed by home gardens, and cultivations

with comparatively low, 18 species (75%) and 10 spe-

cies (42%), respectively. In terms of reptiles, 44 species

(83%), 36 species (68%), and 25 species (47%) were re-

corded in forest edge, home gardens, and cultivations,

respectively (Fig. 11).

Species diversity

Overall the herpetofaunal diversity and both amphibian

and reptile diversity in KPFR was high. The Shannon-

Wiener Index for overall herpetofauna was 3.838.

The Shannon- Wiener Index for amphibian diversity

(7/’^) was 2.508 and for reptile diversity (7/’^) 3.635 (Fig.

12a, b).

Table 1. Checklist of the amphibians {n = 24) recorded from

KPFR. Abbreviations: E - Endemic; EN - Endangered; VU -

Vulnerable; NT - Near Threatened; CE - Closed forest; EE -

Eorest edge; HG - Home Gardens; CU - Cultivations.

Scientific name

Recorded habitats

Ichthyophiidae

Ichthyophis glutinosus ®

Bufonidae

Adenomus kelaartii ®

Duttaphrynus melanostictus

Microhyiidae

Kaloula taprobanica

Micwhyla rubra

Ramanella variegata

Dicrogiossidae

Euphlyctis cyanophlyctis

Euphlyctis hexadactylus

Zakerana kirtisinghei ®

Zakerana syhadrensis

Hoplobatrachus crassus

Nannophrys ceylonensis

Nyctibatrachidae

Lankanectes corrugatus ®

Ranidae

Hylarana aurantiaca

Hylarana temporalis

Rhacophoridae

Pseudophilautus abundus ®

Pseudophilautus cavirostris

Pseudophilautus folicola

Pseudophilautus hoipolloi ®

Pseudophilautus popularis ®

Pseudophilautus reticulatus ®

Pseudophilautus stictomerus

Polypedates cruciger ®

Taruga longinasus

January 2012 | Volume 5 | Number 2 | e39

amphibian-reptile-conservation.org

069

Botejue and Wattavidanage

Table 2. Checklist of the reptiles (n = 53) recorded from KPFR. Abbreviations: E - Endemic; EN - Endangered; VU - Vulnerable;

NT - Near Threatened; CE - Closed forest; EE - Eorest edge; HG - Home Gardens; CU - Cultivations.

Recorded habitats Recorded habitats

Scientific name Scientific name

Pythonidae

Uropeltidae

Python molurus

Rhinophis sp.

Colubridae

Viperidae

Ahaetulla nasuta

Daboia russelii

Ahaetulla pulverulenta

Hypnale hypnale

Amphiesma stolatum

Trimeresurus trigonocephalus ®

Aspidura guentheri

Agamidae

Atretium schistosum

Calotes calotes

Balanophis ceylonensis

Calotes liolepis

Boiga ceylonensis

Calotes versicolor

Boiga forsteni

Ceratophora aspera ™

Cercaspis carinatus

Lyriocephalus scutatus

Chrysopelea ornate

Otocryptis wiegmanni

Coelognathus helena

Gekkonidae

Dendrelaphis bifrenalis

Cnemaspis silvula ®

Dendrelaphis caudolineolatus

Cnemaspis sp.

Lycodon aulicus

Geckoella triedrus

Lycodon osmanhilli ®

Gehyra mutilata

Oligodon arnensis

Hemidactylus depressus ®

Oligodon sublineatus ®

Hemidactylus frenatus

Ptyas mucosa

Hemidactylus parvimaculatus

Sibynophis subpunctatus

Lepidodactylus lugubris ™

Xenochrophis asperrimus ®

Scincidae

Xenochrophis piscator

Eutropis carinata

Cylindrophiidae

Eutropis madaraszi

Cylindrophis maculatus

Lankascincus fallax ®

Elapidae

Lankascincus gansi

Bungarus ceylonicus

Lankascincus greeri ®

Naja naja

Varanidae

Typhlopidae

Varanus bengalensis

Ramphotyphlops sp.

Varanus salvator

Typhlops sp.

Bataguridae

Melanochelys trijuga

Species abundance

During field visits a total of 763 individual amphibians

were recorded, with Zakerana syhadrensis being most

abundant, followed by Euphlyctis cyanophlyctis and E.

hexadactylus. The least abundant species were Ramanel-

la variegata, Pseudophilautus abundus, R cavirostris, P.

reticulatus, and P. stictomerus, followed by Microhyla

rubra, Taruga longinasus, and Ichthyophis glutinosus.

A total of 1,032 individual reptiles were recorded with

Hypnale hypnale being most abundant, followed by

Otocryptis wiegmanni and Lankascincus fallax. The least

abundant species were Ahaetulla pulverulenta, Balano-

phis ceylonensis, Geckoella triedrus, Ramphotyphlops

sp., Typhlops sp., and Rhinophis sp., followed by Aspi-

dura guentheri, Atretium schistosum, Boiga ceylonensis,

and Ceratophora aspera.

Among habitats, abundance was greatest in the for-

est edge, with 269 (35%) individual amphibians and 373

(36%) individual reptiles being recorded. The lowest am-

phibian abundance was documented in closed forest: 158

(20%) individuals; where the lowest reptile abundance

was in cultivations: 171 (17%) individuals. In home gar-

dens, 172 (23%) individual amphibians and 215 (21%)

individual reptiles were recorded, while 164 (22%) indi-

vidual amphibians were recorded in cultivations and 273

(26%) individual reptiles were recorded in closed forest

(Fig. 13).

January 2012 | Volume 5 | Number 2 | e39

amphibian-reptile-conservation.org

070

Herpetofauna of Kalugala proposed forest reserve

Figure 10a. Closely connected cultivation (tea and rubber).

Figure 10b. Closely connected cultivation (tea and paddy).

Habitat type

■ Na. of Amphibian ap. ONa. of RepliFe sp.

Figure 11. Number of species in different habitat types.

Amphibian Reptile Herpetofaunal

diversity diversity diversity

Figure 12a. Herpetofaunal diversity in KPFR.

Closed Forest Forest Edge Home Gardens Cultivations

Habitat type

■ Amphibian Diversity BReptile Diversity

Figure 12b. Herpetofaunal diversity in different habitat types.

40 . 00 %

35 . 00 %

tt 30 . 00 %

^ 25 . 00 %

c 20,00%

^ 15 . 00 %

10 . 00 %

5 . 00 %

0 . 00 %

Closed Forest Forest Edge Home Gardens Cultivations

Habitat type

■ Amphibian abundance ■ Reptile abundance

Figure 13. Species abundance in KPFR.

Species distribution

There were no significant differences in species richness

of amphibians between any habitat type, however, rep-

tiles showed a significant deference in species richness

only between forest edge and cultivations (Mann- Whit-

ney U-test: Z = 2.01, n^ = 44, n^ = 25, P = 0.044).

Discussion

Species richness of amphibians was poor in cultivated

habitats such as tea, rubber, coconut, and some other

commercial crops that are grown in KPFR. However, in

paddy cultivations some dicroglossid frogs were found in

high abundance (e.g., Euphlyctis cyanophlyctis and Za-

kerana syhadrensis). The higher availability of surface

January 2012 | Volume 5 | Number 2 | e39

amphibian-reptile-conservation.org

071

Botejue and Wattavidanage

90 - 00 %

80 . 00 %

70 . 00 %

60 . 00 %

50 . 00 %

40 . 00 %

30 . 00 %

20 . 00 %

10 - 00 %

0 - 00 %

Closed Forest Home Cultivations

Forest Edge Gardens

Habitat type

♦ Prey species ■ Predator species

Figure 14. Distribution of some prey and predator species.

Figure 15. Deforestation inside the KPFR.

water may arguably facilitate these aquatic amphibians

to thrive in paddy cultivations. Euphlyctis cyanophlyc-

tis, however was most abundant in forest edge, along

stream banks and water pools between edges of forest

and cultivations. In home gardens, the most abundant

species were bufonid and dicroglossid frogs including

Duttaphrynus melanostictus, Euphlyctis hexadactylus,

and Zakerana syhadrensis, which is likely related to fa-

vorable living conditions and high abundance of food.

Most of the endemic amphibian species (e.g., Ich-

thyophis glutinosus, Nannophrys ceylonensis, Adenomus

kelaartii, Hylarana temporalis, Pseudophilautus abun-

dus, P. cavirostris, P folicola, P hoipolloi, P popularis,

P reticulatus, P stictomerus, Polypedates cruciger, and

Taruga longinasus) were mostly restricted to the forest

habitats and were commonly not recorded in open areas

such as cultivations and open home gardens. Interest-

ingly, closed forest recorded the lowest amphibian abun-

dance despite having the highest amphibian diversity,

presumably due to high abundance of bufonid and dicro-

glossid frogs in other habitat types.

Figure 16a. Garbage dumping site of the monastery in KPFR.

Figure 16b. Garbage dumping site of the monastery in KPFR.

The distribution pattern of reptile species richness

and species diversity are both similar to amphibians, the

highest being in closed forest and lowest in cultivations.

However, reptile abundance was highest in forest edge

and lowest in cultivations, compared to amphibian abun-

dance, highest in forest edge and lowest in closed for-

est. In cultivations Hypnale hypnale are found in high

numbers potentially, which may be explained by the high

abundance of prey (rodents and frogs) in those cultivat-

ed habitats. Endemic reptile species including Aspidura

guentheri, Balanophis ceylonensis, Cercaspis carinatus,

Dendrelaphis bifrenalis, Xenochrophis asperrimus, Cyl-

indrophis maculatus, Bungarus ceylonicus, Trimeresurus

trigonocephalus, Calotes liolepis, Ceratophora aspera,

Lyriocephalus scutatus, Cnemaspis silvula, Geckoella

triedrus, Hemidactylus depressus, Eutropis madaraszi,

Lankascincus gansi, and L. greeri are mostly forest

dwelling and recorded in lower abundance in other habi-

tats, and rarely in open areas.

Edge effect encompasses biotic and abiotic chang-

es, resulting from the interaction between two different

habitat types (Murcia 1995). Extensive research on edge

effect of many taxa: insects (Hochkirch et al. 2008), am-

phibians (Karunarathna et al. 2008), birds (Helle and

January 2012 | Volume 5 | Number 2 | e39

amphibian-reptile-conservation.org

072

Herpetofauna of Kalugala proposed forest reserve

Helle 1982), and mammals (Pasitschniak-Arts and Mess-

ier 1998). However, Dixo and Martins (2008) show that

edge effects do not influence leaf litter frogs and lizards

in the Brazilian Atlantic forest, despite forest fragmenta-

tion. Similarly, in the present study no edge effects were

detected. The only significant difference among distri-

butions were recorded between forest edge and cultiva-

tions for reptiles (according to Mann-Whitney t/-test).

The forest edge habitats directly adjacent to cultivations

have a high abundance (40%) of reptiles that prey upon

amphibians. In cultivated habitats, dicroglossid and ranid

frogs were found in high abundance possibly due to a

number of water bodies found there (e.g., mud pools and

small rivulets). Therefore, these amphibians may provide

the forage base for the abundant amphibian predatory

reptiles.

Edge effect also applies to succession present where

vegetation is spreading outwards rather than being en-

croached upon. Here, different species are more suited

to edges or central sections of vegetation, resulting in

a varied distribution. In KPFR, many amphibian spe-

cies are normally distributed in higher abundance at

the forest edge rather than other habitats. These include

Ichthyophis glutinosus, Microhyla rubra, Euphlyctis

cyanophlyctis, Zakerana kirtisinghei, Hoplobatrachus

crassus, Lankanectes corrugatus, Hylarana temporalis,

Pseudophilautus abundus, P cavirostris, P folicola, P

hoipolloi, P popularis, P stictomerus, and Taruga lon-

ginasus. Reptiles such as Ahaetulla nasuta, Aspidura

guentheri, Atretium schistosum, Boiga forsteni, Chryso-

pelea ornate, Coelognathus Helena, Dendrelaphis cau-

dolineolatus, Lycodon osmanhilli, Oligodon arnensis,

Sibynophis subpunctatus, Xenochrophis asperrimus, X.

piscator, Cylindrophis maculatus, Bungarus ceylonicus,

Ramphotyphlops sp., Typhlops sp., Rhinophis sp., Calo-

tes calotes, C. liolepis, Otocryptis wiegmanni, Cnemas-

pis silvula, Cnemaspis sp., Hemidactylus depressus, H.

frenatus, H. parvimaculatus, Lepidodactylus lugubris,

Eutropis madaraszi, and Lankascincus greeri have simi-

lar preferences.

The abundance of prey items is much higher than of

predators in all habitats, and predators show distribution

patterns similar to prey, in many instances. For example,

prey species of Euphlyctis and Zakerana show a parallel

distributional pattern to predator species of Xenochro-

phis, Varanus, and Ptyas mucosa (Fig. 14). Species of

Euphlyctis and Zakerana live in a mutual association

(Manamendra-Arachchi and Pethiyagoda 2006) and this

mutual association was clearly observed in KPFR.

Near-primary forest cover accounts for less than

5% of the total wet zone land area, and what remains are

small isolated patches in a sea of human development.

The existing protected forests in the wet zone, which har-

bor a high level of biodiversity, continue to be degraded

due to illegal encroachment and suffer further fragmenta-

tion leading to adverse impacts (lUCNSF and MENRSF

2007).

Adverse human activities have led to deforesta-

tion and habitat loss (Fig. 15) in KPFR. High damage

has been inflicted on the forest habitat by the illegal en-

croachment in forests as a result of improper agriculture

practices and illegal logging; this leads to loss of habitat

and biodiversity. Additionally, the use of agrochemicals

is a great threat to the local biodiversity, especially for the

environmentally sensitive amphibians. Habitual overuse

of agrochemicals in cultivation can lead to death, mal-

formations, and abnormalities in amphibians (de Silva

2009). Most endemic and endangered species found only

in closed forest are at great risk of being exterminated

from the area. One specific threat is the garbage dumps of

the Kalugala Monastery (Fig. 16a, b) which are located

inside the forest.

The material leakage into local streams may worsen

effects on biodiversity as well as the health of people that

inhabit the lower reaches of streams. Material such as

polyethylene bags and other non-biodegradable materi-

als are spread around the monastery and along footpaths

inside the forest. As a result of the garbage dumps, the

population of Varanus salvator and Sus scrofa may have

increased, thus disrupting the ecological balance.

Although these conclusions are based on the results

of this study, we recommend more research be carried out

for longer durations and over a larger area. We strongly

suggest the relevant authorities to take immediate action

to protect this valuable tropical rain forest and to declare

this area a forest reserve, before implementing any long-

term conservation and management plans.

Acknowledgments. — We would like to thank Upali

Amarasinghe (University of Kelaniya, Sri Fanka) for his

help to improve the document and anonymous reviewers

for their valuable comments. The first author wishes to

thank Thasun Amarasinghe and Suranjan Karunarathna

for their valuable comments on the manuscript. We are

grateful to the Conservator General of the Department

of Forest Conservation of Sri Fanka, for allowing the re-

search on herpetofauna in land under their care, and to

the Baduraliya Police for giving assistance to carry out

the held work. We thank Tharindu Sulakshana, Nirmala

Hirantha, Dinesh Gabadage, Suranjan Karunarathna,

Harini Pandithasundara, and Fasanthi Kanthika for as-

sistance during fieldwork and data collection. We thank

I. K. Rajapakshe (The Open University of Sri Fanka), N.

Nilakariyawasam (OUSF), W. C. W. Navaratna (OUSF),

and W. M. P. C. Wijesinghe (OUSF) for all their support.

We also thank the Director General of the Department

of Wildlife Conservation for granting permission to cap-

ture animals for identification; Padmasiri Weerasinghe,

Upathissa Weerasinghe, the villagers of Kalugala for all

their support throughout the study, and Dushantha Kan-

dambi for providing photographs. Finally, we would like

to express our heartfelt gratitude to Ruchira Somaweera,

Kanishka Ukuwela, Craig Hassapakis, and Eric Wild for

the peer-review and editing process.

January 2012 | Volume 5 | Number 2 | e39

amphibian-reptile-conservation.org

073

Botejue and Wattavidanage

Figure 17. Adenomus kelaartii.

Figure 18. Duttaphrynus melanostictus.

Figure 19. Kaloula taprobanica.

Figure 20. Microhyla rubra.

Figure 21. Ramanella variegata.

Figure 22. Euphlyctis hexadactylus.

amphibian-reptile-conservation.org

074

January 2012 | Volume 5 | Number 2 | e39

Herpetofauna of Kalugala proposed forest reserve

Figure 23. Zakerana syhadrensis.

Figure 24. Hoplobatrachus crassus.

Figure 25. Lankanectes corrugatus.

Figure 26. Hylarana aurantiaca.

Figure 27. Pseudophilautus hoipolloi.

Figure 28. Pseudophilautus reticulatus.

January 2012 | Volume 5 | Number 2 | e39

amphibian-reptile-conservation.org

075

Botejue and Wattavidanage

Figure 29. Python molurus.

Figure 31. Atretium schistosum.

Figure 33. Cercaspis carinatus.

Figure 30. Ahaetulla nasuta.

Figure 32. Boiga ceylonensis.

Figure 34. Dendrelaphis caudolineolatus.

Figure 35. Cylindrophis maculatus.

Figure 36. Bungarus ceylonicus.

January 2012 | Volume 5 | Number 2 | e39

amphibian-reptile-conservation.org

076

Herpetofauna of Kalugala proposed forest reserve

Figure 41. Geckoella triedrus.

amphibian-reptile-conservation.org

077

January 2012 | Volume 5 | Number 2 | e39

Botejue and Wattavidanage

Figure 42. Hemidactylus depressus.

Figure 44. Lankascincus greeri.

Figure 46. Varanus bengalensis.

Literature cited

Achard F, Eva HD, Stibig H, Mayaux P, Gallego J, Ricahards T,

Malingreau J. 2002. Determination of deforestation rates of the

world’s humid tropical forests. Science 297(5583):999-1002.

Ashton M, Gunatileke CVS, De Zoysa N, Dassanayake MD, Gu-

NATiLEKE N, WiJESUNDARA S. 1997. A Field Guide to the Common

Trees and Shrubs of Sri Lanka. Wildlife Heritage Trust of Sri

Lanka, Colombo, vii + 432 p.

Bahir mm, Silva a. 2005. Otocryptis nigristigma, a new species of

agamid lizard from Sri Lanka. The Raffles Bulletin of Zoology,

Supplement 12:393-406.

Bahir MM. 2009. Some taxonomic inaccuracies in conservation pub-

lications. Current Science 96(5):632-633.

Figure 43. Eutropis carinata.

Figure 45. Melanochelys trijuga.

Bambaradeniya CNB, Perera MS J, Perera WPN, Wickramasinghe

LJM, Kekulandala LDCB, Samarawickrema VAP, Fernando

RHSS, Samarawickrema VAMPK. 2003. Composition of faunal

species in the Sinharaja world heritage site in Sri Lanka. Sri Lan-

ka Forester 26:21-40.

Batuwita S, Bahir MM 2005. Description of five new species of

Cyrtodactylus (Reptilia: Gekkonidae) from Sri Lanka. The Raffles

Bulletin of Zoology, Supplement 12:351-380.

Batuwita S, Pethiyagoda R. 2007 . Description of a new species of Sri

Lankan litter skink (Squamata: Scincidae: Lankascincus). Ceylon

Journal of Sciences (Biological Sciences) 36(2):80-87.

Bauer AM, de Silva A, Greenbaum E, Jackman T. 2007. Anew spe-

cies of day gecko from high elevation in Sri Lanka, with a prelim-

inary phylogeny of Sri Lankan Cnemaspis (Reptilia, Squamata,

Gekkonidae). Mitteilungen aus dem Zoologischen Museum in

Berlin, Supplement 83:22-32.

Bauer AM, Jackmann TR, Greenbaum E, De Silva A, Giri VB, Das I.

2010a. Molecular evidence for taxonomic status of Hemidactylus

brookii group taxa (Reptilia: Gekkonidae). The Herpetological

Journal 20(3): 129-138.

Bauer AM, Jackman TR, Greenbaum E, Giri VB, De Silva A. 2010b.

South Asia supports a major endemic radiation of Hemidactylus

geckos. Molecular Phylogenetics and Evolution 57(l):343-352.

Bossuyt F, Meegaskumbura M, Beenaerts N, Gower DJ, Pethiya-

goda R, Roelants K, Mannaert A, Wilkinson M, Bahir MM,

Manamendra-Arachchi K, Ng PKL, Schneider CJ, Oomen OV,

Milinkovitch MC. 2004. Local endemism within the Western

Ghats - Sri Lanka Biodiversity Hotspot. Science 306(5695):479-

481.

January 2012 | Volume 5 | Number 2 | e39

amphibian-reptile-conservation.org

078

Herpetofauna of Kalugala proposed forest reserve

Bossuyt F, Meegaskumbura M, Baenerts N, Gower DJ, Pethiya-

GODA R, Roelants K, Mannaert, a, Wilkinson M, Bahir MM,

Manamendra-Arachchi K, Ng PKL, Schneider CJ, Oomen OY,

Milinkovitch MC. 2005. Biodiversity in Sri Lanka and western

Ghats. Science 308(57 19): 199.

Brook BW, Sodhi NS, Ng PKL. 2003. Catastrophic extinctions fol-

low deforestation in Singapore. Nature 424:420-423.

CiNCOTTA RP, W iSNEWSKi J, Engelman R. 2000. Human populations in

the biodiversity hotspots. Nature 404:990-992.

Das I, DE Silva A. 2005. A Photographic Guide to Snakes and Other

Reptiles of Sri Lanka. New Holland Publishers, UK. 144 p.

Dassanayake MD, Fosberg FR (Editors). 1980-1991. A Revised

Handbook of the Flora of Ceylon. Oxford & IBH Publishing Co.,

New Dilhi. Volume 1. vii -i- 508 p.; volume 11. vii -i- 511 p.; volume

111. vii -I- 499 p; volume IV. vii -i- 532 p.; volume V. vii -i- 476 p.;

volume VI. vii -i- 424 p.; volume VIE vii -i- 439 p.

Dassanayake MD, Fosberg FR, Clayon WD (Editors). 1994-1995.

A Revised Handbook of the Flora of Ceylon. Oxford & IBH Pub-

lishing Co., New Dilhi. Volume Vlll. v -i- 458 p.; volume IX. v -i-

482 p.

Dassanayake MD, Clayton WD (Editors). 1996-2000. A Revised

Handbook of the Flora of Ceylon. Oxford & IBH Publishing Co.,

New Dilhi. Volume X. viii -i- 426 p.; volume XL 420 p.; vol. Xll.

390 p.; Xlll. 284 p.; volume XIV. v -i- 307 p.

DE Silva A. 1990. Colour Guide to the Snake Fauna of Sri Lanka. R

and A Publishing Ltd, Avon, England. 130 p.

DE Silva A. 2006. Current status of the reptiles of Sri Lanka In: Fauna

of Sri Lanka: Status of Taxonomy, Research and Conservation.

Editor, Bambaradeniya CNB. The World Conservation Union

(lUCN), Colombo, Sri Lanka and Government of Sri Lanka. 308

p. 134-163.

DE Silva A. 2009. Amphibians of Sri Lanka: A Photographic Guide to

Common Frogs, Toads and Caecilians. Published by the author.

82 plates -i- 168 p.

Deraniyagala pep. 1953. A Colored Atlas of Some Vertebrates from

Ceylon. Volume 2: Tetrapod Reptila. National Museums of Sri

Lanka, Sri Lanka. 101 p.

Deraniyagala PEP. 1955. A Colored Atlas of Some Vertebrates from

Ceylon. Volume 3: Serpentoid Reptila. National Museum of Sri

Lanka, Sri Lanka. 121 p.

Dixo M, Martins M. 2008. Are leaf-litter frogs and lizards affected

by edge effects due to forest fragmentation in Brazilian Atlantic

forest? Journal of Tropical Ecology 24:551-554.

DuttaSK, Manamendra-Arachchi KN. 1996. TheAmphibianFauna

of Sri Lanka. Wildlife Heritage Trust of Sri Lanka, Colombo. 230

P-

F ERNANDO S S, W ICKRAMASINGHA L JM, Rodrigo RK. 2007 . Anew spe-

cies of endemic frog belonging to genus Nannophrys Gunther,

1869 (Anura: Dicroglossinae) from Sri Lanka. Zootaxa 1403:55-

68 .

Frost DR. 2008. Amphibian Species of the World: An Online Refer-

ence. Version 5.2. [Online]. Electronic database. Available: http://

research . amnh . org/herpetology/ amphibia/index .php . [ Acces sed :

15 July 2008]. American Museum of Natural History, New York,

USA.

GunatillekeIAUN, GunatillekeCVS. 1990. Distribution offloristic

richness and its conservation in Sri Lanka. Conservation Biology

4(1):21-31.

Gunther ALCG. 1864. The Reptiles of British India. Ray Society,

Robert Hardwicke, London, xxii -i- 26 plates -i- 452 p.

Gower DJ, Maduwage K. 2011. Two new species of Rhinophis

Hemprich (Serpentes: Uropeltidae) from Sri Lanka. Zootaxa

2881:51-68.

Helgen km. Groves CP. 2005. Biodiversity in Sri Lanka and western

Ghats. Science 308(5719): 199.

Helle E, Helle P 1982. Edge effect on forest bird densities on off-

shore islands in the northern Gulf of Bothnia. Annales Zoologici

Fennici 19:165-169.

Hewawasam T, von Blanckenburg F, Schaller M, Kubik P. 2003.

Increase of human over natural erosion rates in tropical highlands

constrained by cosmogenic nuclides. Geology 31:597-600.

Hochkirch A, Gartner A-C, Brandt T. 2008. Effects of forest-dune

forest edge management on the endangered heath grasshopper,

Chorthippus vagans (Orthoptera: Acrididae). Bulletin of Entomo-

logical Research 98:449-456.

Howlader MSA. 2011. Cricket frog (Amphibia: Anura: Dicroglos-

sidae): two regions of Asia are Corresponding two groups. Bon-

noprani - Bangladesh Wildlife Bulletin 5(1-2): 1-7.

Jenkins M. 2003. Prospects for biodiversity. Science 302(5648): 1175-

1177.

KarunarathnaDMSS,AbeywardanaUT1,AselaMDC,Kekuland-

ALA EDCB. 2008. A preliminary survey of the amphibian fauna in

Nilgala forest area and its vicinity, Monaragala district, Sri Lanka.

Herpetological Conservation and Biology 3(2):264-272.

Kekulandala LDCB. 2002. A survey of fish in Kalugala Proposed

Forest Reserve. Sri Lanka. Naturalist 5(2):20-25.

Maduwage K, Silva A, Manamendra-Arachchi K, Pethiyagoda R.

2009. A taxonomic revision of the South Asian hump-nosed pit

vipers (Squamata: Viperidae: Hypnale). Zootaxa 2232:1-28.

Manamendra-Arachchi K, Pethiyagoda R. 2005. The Sri Lankan

shrub-frogs of the genus Philautus Gistel, 1848 (Ranidae: Rha-

cophorinae) with description of 27 new species. The Raffles Bul-

letin of Zoology, Supplement 12:163-303.

Manamendra-Arachchi K, Pethiyagoda R. 2006. Sri Lankawe Ub-

hayajeeween [Amphibians of Sri Lanka]. Wildlife Heritage Trust

of Sri Lanka, Colombo. 88 plates -i- 440 p.

Manamendra-Arachchi K, de Silva A, Amarasinghe T. 2006. De-

scription of a second species of Cophotis (Reptilia:Agamidae)

from the highlands of Sri Lanka. Lyriocephalus, Supplement

l(6):l-8.

Manamendra-Arachchi K, Batuwita S, Pethiyagoda R. 2007. A

taxonomic revision of the Sri Lankan day-geckos (Reptilia: Gek-

konidae: Cnemaspis), with description of new species from Sri

Lanka and southern India. Zeylanica 7(1):9-122.

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, Sup-

plement 12:305-338.

Meegaskumbura M, Manamendra-Arachchi K. 20 1 1 . Two new spe-

cies of shrub frogs (Rhacophoridae: Pseudophilautus) from Sri

Lanka. Zootaxa 2747:1-18.

January 2012 | Volume 5 | Number 2 | e39

amphibian-reptile-conservation.org

079

Botejue and Wattavidanage

Meegaskumbura M, Bossuyt F, Pethiyagoda R, Manamendra-

ArachchiK, BahirMM, MilinkovitchMC, SchneiderCJ. 2002.

Sri Lanka: An amphibian hotspot. Science 298(5592):379.

Meegaskumbura M, Manamendra-Arachchi K, Pethiyagoda R.

2009. Two new species of shrub frogs (Rhacophoridae: Philautus)

from the lowlands of Sri Lanka. Zootaxa 2122:51-68.

MeegaskumburaM,Manamendra-ArachchiK,SchneiderCJ,Pethi-

YAGODA R. 2007. New species amongst Sri Lanka’s extinct shrub

frogs (Amphibia: Rhacophoridae: Philautus). Zootaxa 1397:1-15.

Meegaskumbura M, Meegaskumbura S, Bo watte G, Manamendra-

Arachchi K, Pethiyagoda R,HankenJ, SchneiderCJ. 2010. Ta-

ruga (Anura: Rhacophoridae), a new genus of foam-nesting tree

frogs endemic to Sri Lanka. Ceylon Journal of Science (Biologi-

cal Sciences) 39(2):75-94.

Murcia C. 1995. Edge effects in fragmented forest: Implication for

conservation. Trends in Ecology and Evolution 10(2):58-62.

Pasitschniak-Arts M and Messier F. 1998. Effects of edges and habi-

tats on small mammals in a prairie ecosystem. Canadian Journal

of Zoology 76(ll):2020-2025.

Pethiyagoda R. 1994. Threats to the indigenous freshwater fishes

of Sri Eanka and remarks on their conservation. Hydrobiologia

285(1-3):189-201.

Pethiyagoda R. 2005. Exploring Sri Eanka’ s biodiversity. The Raffles

Bulletin of Zoology, Supplement 12:1-4.

Pethiyagoda R. 2001 di. Pearls, Spices and Green Gold, An Illustrated

History of Biodiversity Exploration of Sri Lanka. Wildlife Heri-

tage Trust of Sri Eanka, Colombo. 241 p.

Pethiyagoda R. 2007b. The ‘New species syndrome’ in Sri Eankan

herpetology: A cautionary note. Zeylanica 7(1): 1-7.

Pethiyagoda R, Manamendra-Arachchi K. 1998. A revision of the

endemic Sri Eankan agamid lizard genus Ceratophora Gray,

1835, with description of two new species. Journal of South Asian

Natural History l(l):77-96.

Ranasinghe PN. 1 995 . Biodiversity and Management Plan: Kalugala

Proposed Eorest Reserve. The Young Zoologists’ Association,

National Zoological Gardens, Dehiwala. 45 p. (Unpublished).

Senaratna EK. 2001. A Check List of the Elowering Plants of Sri

Lanka. National Science Foundation of Sri Eanka, Colombo. 342

P-

Smith EN, Manamendra-Arachchi K, Somaweera R. 2008. Anew

species of coral snake of the genus Calliophis (Squamata: Elapi-

dae) from the Central Province of Sri Eanka. Zootaxa 1 847 : 1 9-33 .

Smith MA. 1935. The Eauna of British India Including Ceylon and

Burma: Reptilia and Amphibia, Volume II Sauria. Taylor and

Francis, Eondon. 1 plate -i- 440 p.

Somaweera R. 2006. Sri Lankawe Sarpayin [Snakes of Sri Eanka].

Wildlife Heritage Trust of Sri Eanka, Colombo. 297 p.

Somaweera R and Somaweera N. 2009. Lizards of Sri Lanka: A Co-

lour Guide with Eield Keys. Edition Chimaira, Serpent’s Tail,

German. 303 p.

Tan BC. 2005. New species records of Sri Eanka Mosses. The Raffles

Bulletin of Zoology, Supplement 12:5-8.

Taylor EH. 1953. A review of the lizards of Ceylon. The University

of Kansas Science Bulletin 35(12): 1525-1585.

lUCN Sri Lanka (IUCNSL), The Ministry of Environment and Natu-

ral Resources (MENRSE). 2007. The 2007 Red List of threatened

Eauna and Elora of Sri Lanka. The World Conservation Union

(lUCN) Colombo, Sri Eanka. 148 p.

Whitaker R and Captain A. 2004. Snakes of India, the Eield Guide.

Draco Publication Eimited, India. 479 p.

WickramasingheLJM, MunindradasaDAI. 2007. Review ofthe ge-

nus Cnemaspis Strauch, 1887 (Sauria: Gekkonidae) in Sri Eanka

with the description of five new species. Zootaxa 1490: 1-63.

WiCKRAMASINGHE LJM, ViDANAPATHIRANA DR, WiCKRAMASINGHE N,

Ranwella PN. 2009. A new species of Rhinophis Hemprich, 1 820

(Reptilia: Serpentes: Uropeltidae) from Rakwana massif, Sri Ean-

ka. Zootaxa 2044:1-22.

WiTTEMYER G, Elsen P, Bean WT, Burton ACO, Brashares JS . 2008.

Accelerated human population growth at protected area edges.

Science 321(5885): 123-126.

Manuscript received: 16 November 2011

Accepted: 10 December 2011

Published: 2 February 2012

W. MADHAVA S. BOTEJUE has

researched the fauna of Sri Lanka

for the past seven years, especially

ecology and behavior. He has con-

ducted awareness programs to ed-

ucate the Sri Lankan community of

the importance of biodiversity and

conservation. Madhava earned his

B.Sc. degree in Natural Sciences

from The Open University of Sri

Lanka (OUSL) in 2009. Currently

he serves as treasurer of the Taprobanica Nature Conservation

Society, Sri Lanka and associate editor for Taprobanica: The

Journal of Asian Biodiversity.

DR. JAYANTHA WATTAVI-

DANAGE has been involved in

teaching and research in the fields

of ecology, faunal diversity, lim-

nology, and molecular parasitolo-

gy for the past twenty years. He is

strongly involved in popularizing

science among the general public

and is the author of a large num-

ber of newspaper and magazine

articles. Currently, he is the Chair-

man, National Committee for Science Popularizing, National

Science Eoundation in Sri Lanka. He works as a Senior Lectur-

er in zoology at The Open University of Sri Lanka beginning in

1990. Jayantha earned his B.Sc. and M.Phil. from University of

Sri Jayawardenepura and Ph.D. from University of Colombo,

Sri Lanka. He conducted post doctoral research on molecular

genetics of Malaria at University of Edinburgh, United King-

dom. He is also a recipient of many research awards including

the Presidential Award for his research publications.

January 2012 | Volume 5 | Number 2 | e39

amphibian-reptile-conservation.org

080

Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium,

provided the original author and source are credited.

Amphibian & Reptiie Conservation 5(2):81-89.

Herpetofauna in the Kaluganga upper catchment of the

Knuckles Forest Reserve, Sri Lanka

1 ^V.A.M.P.K. SAMARAWICKRAMA, ^D.R.N.S. SAMARAWICKRAMA, AND ^SHALIKA KUMBUREGAMA

^No:308/7A, Warathenna, Halloluwa, Kandy, SRI LANKA ^Department of Zoology, University of Peradeniya, SRI LANKA

Abstract . — ^The Knuckles Forest Reserve and forest range is a paradise for a large number of endem-

ic Sri Lankan taxa, including a considerable number of amphibian and reptile species. A survey car-

ried out on the western slopes of the Kaluganga catchment of Knuckles Forest Reserve recorded 19

species of amphibians and 30 species of reptiles. Of these, 15 species of amphibians and 17 species

of reptiles are endemic to Sri Lanka, and 11 species are restricted to a few localities in the Knuckles

forest range. Three unidentified species possibly new to science were discovered in the study, and

we recommend that these species need further study for taxonomic identification.

Key words. Knuckles forest reserve, herpetofauna, endemic, restricted, threatened, Sri Lanka

Citation: Samarawickrama VAMPK, Samarawickrama DRNS, Kumburegama S. 2012. Herpetofauna in the Kaluganga upper catchment of the Knuckles

Forest Reserve, Sri Lanka. Amphibian & Reptile Conservation 5{2)■.8^ -89 (e41).

Introduction

The Knuckles mountain range of Sri Lanka is a distinct

topographic feature of the central highlands of Sri Lanka,

covering approximately 21,000 ha. It lies between lati-

tudes 7°18’-7°34’ N and longitudes 80°41’-80°55’ E at

900-1900 m elevation range. This landscape is made

unique by the aggregation of at least 35 spectacular peaks

rising above 900 m in the Kandy and Matale Districts.

The Knuckles range is geologically part of the central

highlands of the island but isolated from the main moun-

tain mass by the Mahaweli River valley on the south and

east and on the west by the Matale valley (De Rosayro

1958). The Knuckles range is one of the more important

watersheds in the country. It receives rainfall from both

the southwest and northeast monsoons. Numerous tribu-

taries of the Knuckles contribute to major rivers, includ-

ing the Mahaweli. The area’s mean annual temperature

outside the massif is more than 26 °C, and this value falls

to about 21 °C at elevations above 915 m and to about

18.5 °C at the highest elevations (Cooray 1998).

The topographic and climatic variation in the

Knuckles region has resulted in the occurrence of sev-

eral natural vegetation types. According to Rosayro

(1958), vegetation types of the Knuckles region are cat-

egorized as lowland tropical wet semi-evergreen forests,

sub-montane tropical wet semi-evergreen forests, and

montane tropical wet evergreen forests. Gunatilleke and

Gunatilleke (1990) recognized 15 floristic regions in Sri

Lanka, and each of these has dominant plant communi-

Correspondence. Email: ^madurapk@yahoo.com

ties. The Knuckles forest belongs to the 12* floristic re-

gion (termed Knuckles) with a unique vegetation type.

According to these authors, there are two types of natural

vegetation in this region: tropical montane forests charac-

terized by a Calophyllum zone and tropical sub-montane

forests characterized by a Myristica, Cullenia, Aglaia,

and Litsea community (Karunarathna et al. 2009).

In addition to these categories, there are anthropo-

genic vegetation types such as patana grasslands, which

are dominated by Cymbopogon spp. derived from aban-

doned coffee and tea plantations, scrublands, and agri-

cultural land.

The geographic location, altitude, and position of the

mountain range in relation to the two main wind currents

that cross the island have resulted in a unique ecosystem

with an abundance of endemic flora and fauna (Kariya-

wasam 1991). The variety of habitats and forest com-

munities in the Knuckles is known to harbor a diverse

community of herpetofauna, but a large extent of the

mountain range remains unexplored. In an effort to iden-

tify and study the distribution of amphibians and reptiles,

a study was carried out in the tropical montane forests,

sub-montane forests, and lowland semi-evergreen for-

ests of the under-researched Kaluganga catchment of the

Knuckles range. These forest types were derived based

on elevational range (Bambaradeniya and Ekanayake

2003):

• Tropical Montane Forest (>1300 m a.s.l.)

• Tropical Sub-montane Forest (600-1300 m a.s.l.)

• Lowland Semi-evergreen Forest (below 700 m a.s.l.)

amphibian-reptile-conservation.org

081

March 2012 I Volume 5 I Number 2 I e41

Samarawickrama et al.

Tropical Sub-montane Forest (600-1300 m a.s.l.).

Lowland Semi-evergreen Forest (below 700 m a.s.l).

Tropical Montane Forest (>1300 m a.s.l).

Methods

Fieldwork was conducted from May to July 2010 in the

Kaluganga upper catchment of Knuckles range. The

study area extended from the Pallegama main bridge to

Kalupahana mountain area. In each habitat, data were

collected from five 100 x 10 m transects, with one night

sampling per habitat. The distance between transects

was more than 500 m. Within each major habitat, dif-

ferent microhabitats (such as tree trunks, tree holes, wa-

ter puddles, and other small niches) were systematically

searched for herpetofauna. Three people were involved

in the sampling of each transect. One person searched

above 1.5 m on trees for arboreal species, while a second

person pursued a terrestrial search under logs, stones, leaf

litter, tree trunks, etc., and a third person searched aquatic

habitats (puddles and streams). In addition to recording

the different species within each transect, a thorough

search for different amphibians and reptiles was carried

out along nature trails or footpaths and streams outside

of the five transects. The different species of amphibians

and reptiles were hand-captured or collected using a hand

net and observed. Frog species were located using their

call signatures. Taxonomic keys (Manamendra-Arachchi

and Pethiyagoda 2006; Dutta and Manamendra-Arachchi

1996; De Silva 1980; Deraniyagala 1953; Somaweera

2006; Taylor 1953) were used for identification or con-

firmation of collected species. Photographs of live speci-

mens were taken in the field using a Canon EOS 350 SLR

camera. After identification, the animals were released to

their natural habitat unharmed.

Results and discussion

A total of 49 species of amphibians and reptiles were

identified from the study sites. The survey documented

19 species of amphibians belonging to the families Bu-

fonidae, Dicroglossidae, Nyctibatrachidae, Ranidae, and

Amphib. Reptile Conserv. | http://redlist-ARC.org 082

March 2012 I Volume 5 | Number 2 | e41

Herpetofauna in the Kaluganga, Knuckles Forest Reserve, Sri Lanka

Table 1. List of amphibians recorded during the study period from the Kaluganga upper catchment in the Knuckles

(Abbreviations: * - Endemic to Sri Lanka; /R - restricted to the Knuckles forest region; CR - Critically Endangered; and

EN - Endangered).

Family

Scientific name

Common name

Bufonidae

Adenomus kelaartii *

Kelaart’s dwarf toad

Duttaphrynus melanostictus

Common house toad

Dicroglossidae

Euphlyctis cyanophlyctis

Skipper frog

Fejervarya kirtisinghei *

Mountain paddy field frog

Fejervarya limnocharis

Common paddy field frog

Nannophrys marmorata

Kirtisinghe’s rook frog

Nyctibatrachidae

Lankanectes cf. corrugatus *

Corrugated water frog

Ranidae

Hylarana temporalis

Common wood frog

Hylarana gracilis *

Sri Lanka wood frog

Rhacophoridae

Pseudophilautus fergusonianus *

Ferguson’s tree frog

Pseudophilautus fulvus

Knuokles shrub frog

Pseudophilautus hoffmanni *’

Hoffmann’s shrub frog

Pseudophilautus hankeni *’*

Hanken’s shrub frog

Pseudophilautus stuarti *’

Stuart's shrub frog

Pseudophilautus steineri

Steiner’s shrub frog

Pseudophilautus macropus

Bigfoot shrub frog

Pseudophilautus cavirostris

Tuberole tree frog

Polypedates cruciger *

Common hour-glass tree frog

Taruga cf. eques

Mountain hourglass tree frog

Rhacophoridae (15 of these species are endemic to the

island; Table 1). In addition, three unidentified species of

amphibians were collected; further studies are being car-

ried out for taxonomic identification of these three spe-

cies, and they may or may not be new to science. Further

studies are also being carried out to identify the distribu-

tion and ecology of Taruga eques and Lankanectes cf.

corrugatus in the region.

Among the identified species, there are seven re-

gionally endemic species restricted to the Knuckles

range, including three Critically Endangered species

(Pseudophilautus hankeni, R macropus, and Nannoph-

rys marmorata) and six Endangered species (P. fulvus,

R hoffmanni, R stuarti, R steineri, R cavirostris, and Ta-

ruga eques).

In this study, a total of 30 species of reptiles were

recorded, with 17 regionally endemic species including

four species restricted to Knuckles (Table 2). Among

Adenomas kelaartii.

these, two species are Critically Endangered {Cophotis

dumbara and Chalcidoseps thwaitesi) and four species

are Endangered (Calotes liocephalus, Ceratophora ten-

nentii, Cyrtodactylus soba, and Lankascincus deraniya-

galae) (lUCN-SE and MENR-SE 2007).

Brief description of naturai history and dis-

tribution of key species encountered during

survey

Adenomus kelaartii

Endemic species to the island and found in lowland semi-

evergreen forests of Knuckles forest range, primarily in

riverine forests and wet patana grasslands. Species com-

monly observed on leaf litter and rarely recorded in semi-

arboreal habitats 1.5 m above ground. Species recorded

from Rambukoluwa and Manigala patana area.

Nannophrys marmorata.

Amphib. Reptile Conserv. | http://redlist-ARC.org 083

March 2012 I Volume 5 | Number 2 | e41

Samarawickrama et al.

Table 2. Reptiles recorded during study period from Kaluganga upper catchment Knuckles range (Abbreviations:

Endemic to Sri Lanka; /R - restricted to the Knuckles forest region; CR - Critically Endangered; and EN - Endan-

gered).

Family

Scientific name

Common name

Agamidae

Calotes calotes

Green garden lizard

Calotes liolepis *

Whistling lizard/Forest lizard

Calotes liocephalus

Crestless lizard

Calotes versicolor

Common garden lizard

Cophotis dumbara *’

Dumbara pigmy lizard

Ceratophora tennentii

Leaf nose lizard

Lyriocephalus scutatus *

Lyre-head lizard/Hump snout lizard

Otocryptis wiegmanni *

Sri Lankan kangaroo lizard

Gekkonidae

Cnemaspis kallima *

Ornate day geeko

Cyrtodactylus soba *®’En

Knuekles forest geeko

Gehyra mutilata

Four-claw gecko

Hemidactylus parvimaculatus

Spotted house geeko

Hemidactylus depressus *

Kandyan geeko

Hemidactylus frenatus

Common house-geeko

Scincidae

Dasia haliana *

Haly’s tree skink

Lankascincus deraniyagalae *’

Deraniyagala's lanka skink

Lankascincus taprobanensis *

Smooth lanka skink

Mabuya macularia

Bronze-green little skink

Chalcidoseps thwaitesii *’

Four- toe snake skink

Colubridae

Ahaetulla nasuta

Green vine snake

Ahaetulla pulverulenta

Brown vine snake

Boiga ceylonensis

Sri Lanka eat snake

Dendrelaphis caudolineolatus

Gunther’s bronze baek

Dendrelaphis tristis

Common bronze baek

Macropisthodon plumbicolor

Green keelbaek

Oligodon sublineatus *

Dumerul’s kuki snake

Ptyas mucosa

Rat snake

Elapidae

Calliophis haematoetron *

Blood-bellied eoral snake

Viperidae

Hypnale cf. nepa *

Merrem’s hump-nosed viper

Trimeresurus trigonocephalus *

Green pit viper

Lankanectes cf. corrugatus

Lankanectes is a monotypic genus. Endemic species

commonly found in the wet zone. Our data suggest the

Lankanectes sp. observed in Knuckles is distinct from

L. corrugatus found elsewhere; a taxonomic study is be-

ing carried out to understand its relationship within the

genus. Recorded from montane and sub-montane forest

habitats and commonly found in rocky-bottomed streams

and water holes.

Nannophrys marmorata

Endemic, Critically Endangered species restricted to

the Knuckles. Only recorded in Patana grasslands found

within sub-montane and lowland semi-evergreen forests

and in moist rock crevices. There are two other species

recorded in this genus: N. ceylonensis found in the low-

land wet zone and N. naeyakai restricted to the Uva and

eastern provinces of Sri Lanka (Fernando et al. 2007).

Pseudophilautus cavirostris

Endemic, Endangered species recorded from lowland

semi-evergreen forests and Kaluganga riverine forests

on tree trunks about L5-2 m above ground. Prefers to

remain under thick, moist moss on tree trunks. Primar-

ily found from Pallegama to Rambukoluwa (Kaluganga

river bank).

Amphib. Reptile Conserv. | http://redlist-ARC.org 084

March 2012 I Volume 5 | Number 2 | e41

Herpetofauna in the Kaluganga, Knuckles Forest Reserve, Sri Lanka

Pseudophilautus cavirostris.

Pseudophilautus hankeni.

Pseudophilautus fergusonianus.

Pseudophilautus macropus.

Pseudophilautus fergusonianus

Pseudophilautus fergusonianus was recorded from low-

land semi-evergreen forests in the study area. Endemic

species primarily found on moist rock surfaces near

streams during the day and on shrubs at night. Recorded

in Walpalamulla and Rambukoluwa area.

Pseudophilautus fulvus

Endemic and Endangered species primarily found in sub-

montane and lowland semi-evergreen forests. They occu-

py small tree holes during day and at night were observed

on tree bark. Species recorded from Bambarakanda (near

Walpalamulla). Only a single specimen was documented

in this study.

Pseudophilautus hankeni

Psuedophilautus hankeni a recently described species

(Meegaskumbura and Manamendra-Arachchi 2011);

conservation status not assessed yet. Species only re-

corded from the Knuckles range and was previously re-

corded only in Dothalugala Man and Biosphere Reserve

within the Knuckles conservation forest (Rajapaksha et

al. 2006). Uncommon, arboreal species. Major habitat

is montane forests living on mossy tree bark; occasion-

ally recorded on ground. Documented from Kalupahana

mountain range, Gomabaniya, and Yakungehela areas,

expanding its previous range.

Pseudophilautus macropus

Endemic, Critically Endangered amphibian primarily

found near streams in sub-montane forest habitats. Only

one specimen was recorded during the study, collected

on mossy bark, about 1.5 m from the ground in the Bam-

barakanda area.

Pseudophilautus stuarti

Endemic and Endangered species restricted to the Knuck-

les forest range found in understory of montane and sub-

montane forest habitats, mostly in shrub layer. Recorded

in Kalupahana peak, Gombaniya northern slope, and

Bambarakanda.

Hylarana gracilis

Endemic species primarily recorded from riverbanks of

lowland semi-evergreen forests. Ground-living species,

recorded from banks of the Kaluganga Pallegama to

Rambukoluwa rivers.

Amphib. Reptile Conserv. | http://redlist-ARC.org 085

March 2012 I Volume 5 | Number 2 | e41

Samarawickrama et al.

Pseudophilautus stuarti.

Calotes liocephalus.

Calotes liolepis.

Cophotis dumbara.

Calotes liocephalus

Endemic, Endangered, and arboreal, found in the Kalu-

pahana peaks. Only a single specimen was documented

in this study.

Calotes liolepis

Endemic arboreal species found on tree branches four m

above ground in the Walpallamulla area. Agile and fast-

moving.

Cophotis dumbara

Endemic and Critically Endangered species recorded

from outside of the transect. Restricted to the Knuckles

range; there are only a few records of this enigmatic spe-

cies. First documentation of this species from Kalupa-

hana mountain area. Only one specimen was recorded

basking 1.5 m above ground.

Ceratophora tennentii

Ceratophora tennentii is an endemic. Endangered spe-

cies, restricted to the Knuckles range. Species found in

montane and sub-montane forest habitats. Semi-arboreal,

found both on and above ground. Species recorded from

Kalupahana, Bambaragala, and Gombaniya peaks.

Lyriocephalus scutatus

Endemic species with its major habitat in lowland semi-

evergreen forests. Species found 1.5 m above ground,

close to Yakungehela area. Display of deep red color is a

defensive behavior in this species.

Cyrtodactylus soba

Endemic and Endangered species restricted to the Knuck-

les forest. Species recorded in montane forest habitats

and rock crevices in Yakungehela peaks.

Chalcidoseps thwaitesi

Endemic and Critically Endangered species only previ-

ously recorded in a few localities in Knuckles range in

lowland semi-evergreen forests. Fossorial species found

under rocks in Yakungehela area.

Dasia halianus

Endemic species (Wickramasinghe et al. 2011) observed

basking on tree bark in lowland semi-evergreen forests

near Rambukoluwa area.

Amphib. Reptile Conserv. | http://redlist-ARC.org 086

March 2012 I Volume 5 | Number 2 | e41

Herpetofauna in the Kaluganga, Knuckles Forest Reserve, Sri Lanka

Ceratophora tennentii.

Cyrtodactylus soba.

Lyriocephalus scutatus.

Dasia halianus.

Calliophis haematoetron.

Calliophis haematoetron

Conclusion

Endemic, recently described species (Smith et al. 2008),

and one of two species of coral snakes found in the coun-

try. Species recorded only from a few localities and Pal-

legama semi-evergreen forest. Fossorial form found on

thick leaf litter layers.

Trimeresurus trigonocephalus

Endemic species exhibiting different color morphs and

found in lowland semi-evergreen and sub-montane for-

ests (plain green variation found). Nocturnal species,

mostly found on bushes and in tree holes.

This survey is indicative of the importance of Knuckles

range in providing refuge to a large number of amphibian

and reptile species. These species are facing habitat loss,

mainly due to anthropogenic activities. Forest encroach-

ment, seasonal fires on the dry phase of the Knuckles

range, illegal felling of trees, occasional gem mining, and

cardamom plantations are among the threats faced by the

diverse species in the Knuckles. Over several decades,

the forests in the Knuckles have degraded due to carda-

mom planting, and to a lesser extent, by shifting culti-

vation and potato growing (Kariyawasam 1991). Carda-

mom plants thrive in shady, cool, and humid conditions

at high elevations, so cardamom planters remove part of

Amphib. Reptile Conserv. | http://redlist-ARC.org 087

March 2012 I Volume 5 | Number 2 | e41

Samarawickrama et al.

Chalcidoseps thwaitesi.

the canopy and clear understory of the forest. These ac-

tivities may be extremely detrimental to some species. In

addition, similar to what is observed in the Horton Plains

National Park in Sri Lanka, forest dieback also occurs

in large tracts of forest in the Knuckles range. Causes of

this dieback are uncertain. The resulting forest destruc-

tion and fragmentation will certainly have an adverse

effect on its inhabitants. Herpetofauna in particular are

extremely vulnerable to habitat changes (Pierce 1985;

Wyman 1990; Blaustein et al. 1998). Furthermore, habi-

tat loss and fragmentation due to any number of reasons

will be especially detrimental to species restricted to the

Knuckles. Further studies and strict conservation mea-

sures are necessary to help safeguard the herpetofauna

and all the flora and fauna, that are maintaining a delicate

balance in this ecosystem.

Acknowledgments. — The authors thank Anura Ban-

dara and Prasad Wijesekara for their valuable held as-

sistance. Also, we would like to thank Craig Hassapakis,

Rohan Pethiyagoda, Suranjan Karunarathna, and anony-

mous reviewers who helped in diverse ways to enrich

this work.

Literature cited

Bambaradeniya CNB, Ekanayake SP. 2003. A Guide to

the Biodiversity of Knuckles Forest Region. lUCN,

Trimeresurus trigonocephalus.

Colombo, Sri Lanka. 68 p.

Blaustein AR, Kiesecker JM, Chivers DP, Hokit DG,

Marco A, Belden LK, Hatch A. 1998. Effects of

ultraviolet light on amphibians: Field experiments.

American Zoologist 38(6):799-812.

ChandrajithR, KoralegedaraN, RanawanaKB, Tob-

SCHALL HJ, Dissanayake CB. 2009. Major and trace

elements in plants and soils in Horton Plains National

Park, Sri Lanka: An approach to explain forest die

back. Environmental Geology 57(1): 17-28.

CooRAY PG. 1998. The Knuckles Massif. A Portfolio, For-

est Department, Battaramulla, Sri Lanka. 76 p.

De Rosayro RA. 1958. The climate and vegetation of the

Knuckles region of Ceylon. The Ceylon Forester 3(3-

4):210-260.

Deraniyagala pep. 1953. A Colored Atlas of Some Ver-

tebrates from Ceylon. Tetrapod Reptila Volume 2.

Ceylon National Museums publication, Colombo, Sri

Lanka. 101 p.

De Silva PHDH. 1980. Snake Fauna of Sri Lanka with

Special Reference to Skull, Dentition and Venom in

Snakes. National Museums of Sri Lanka, Colombo,

Sri Lanka. 472 p.

Dutta SK, Manamendra-Arachchi K. 1996. The Am-

phibian Fauna of Sri Lanka. The Wildlife Heritage

Trust, Colombo, Sri Lanka. 230 p.

Fernando SS, Wickramasingha LJM, Rodirigo RK.

2007. A new species of endemic frog belonging to

genus Nannophrys Gunther, 1869 (Anura: Dicroglos-

sinae) from Sri Lanka. Zootaxa 1403:55-68.

Gunatilleke IAUN, Gunatilleke CVS. 1990. Distribu-

tion of floristic richness and its conservation in Sri

Lanka. Conservation Biology 4(1):21-31.

lUCN-SL, MENR-SL. 2007. The 2007 List of Threat-

ened Fauna and Flora of Sri Lanka. lUCN Sri Lanka,

Colombo, Sri Lanka, xiii + 148 p.

Kariyawasam D. 1991. Resource use and settlement in

the forests of the Knuckles Range. The Sri Lanka For-

ester 20 (New series; 1&2):3-13.

Karunarathna, DMSS, Bandara IN, Chanaka AWA.

2009. The ovipositional behavior of the endemic

Amphib. Reptile Conserv. | http://redlist-ARC.org 088 March 2012 | Volume 5 | Number 2 | e41

Herpetofauna in the Kaluganga, Knuckles Forest Reserve, Sri Lanka

lizard Calotes liolepis Boulenger, 1885 (Reptilia:

Agamidae) in the Knuckles forest region of Sri Lanka.

Acta Herpetologica 4(l):47-56.

Manamendra-Arachchi K, Pethiyagoda R. 2006. Sri

Lankawe Ubaya jeeween (Amphibians of Sri Lanka)

Colombo, Sri Lanka. 440 p. (Text in Sinhala).

Meegaskumbura M, Manamendra-Arachchi K. 2011.

Two new species of shrub frogs (Rhacophoridae:

Pseudophilautus) from Sri Lanka. Zootaxa 2747:1-

18.

Pierce BA. 1985. Acid tolerance in amphibians. BioSci-

ence 35(4):239-243.

RajapakshaDRNS, Samarawickrma VAMPK, Ranawa-

NA KB. 2006. Herpetofaunal diversity in Dothalugala

MAB Reserve, of the Knuckles forest range, Sri Lan-

ka. Russian Journal of Herpetology 13(2): 120-134.

Smith EN, Manamendra-Arachchi K, Somaweera R.

2008. A new species of coralsnake of the genus Cal-

liophis (Squamata: Elaphidae) from the Central Prov-

ince of Sri Lanka. Zootaxa 1847:19-33.

Somaweera R. 2006. Sri Lankawe Sarpayan (The Snakes

of Sri Lanka, Colombo, Sri Lanka. 270 p. (Text in Sin-

hala).

Taylor EH. 1953. A review of the lizards of Ceylon.

University of Kansas Science Bulletin 35 (Part 11, no.

12):1525-1585.

WiCKRAMASINGHE EJM, WiCKRAMASINGHE N, KaRIYA-

WASAM L. 2011. Taxonomic status of the arboreal

Skink Lizard Dasia halianus (Haly & Nevill, 1887) in

Sri Lanka and the redescription of Dasia subcaerule-

um (Boulenger, 1891) from India. Journal of Threat-

ened Taxa 3(8): 1961-1974.

Wyman RL. 1990. What’s happening to the amphibians?

Conservation Biology 4(4): 350-352.

Manuscript received: 10 November 2011

Accepted: 04 December 2011

Published: 02 March 2012

V.A.M.P.K. SAMARAWICKRAMA has about fifteen years field experience with herpetofauna and

worked as an ecologist for the lUCN Sri Lanka County Office. His research over the past decade has

been on taxonomical studies of herpetofauna, animal behavioral studies, and he has described a new

species of lizard from the Knuckles forest region; his current research concerns the genus Lankanectes.

Hobbies are nature painting, outdoor camping, bird watching, wildlife photography, and hiking.

D.R.N.S. SAMARAWICKRAMA graduated from the University of Peradeniya in 2001 and obtained

her master’s degree (M.Sc.) in Environmental Forestry from the Post Graduate Institute of Agriculture,

University of Peradeniya. Her research project was titled, “Herpetofaunal Diversity in Dothalugala

MAB Reserve, of the Knuckles Forest Range, Sri Lanka.” Her hobbies include hiking and outdoor

camping.

SHALIKA KUMBUREGAMA was trained as a developmental biologist and obtained her Ph.D. in

Zoology from University of Hawaii at Manoa in 2009. Currently, she is working as a Senior Lecturer

in the Department of Zoology, University of Peradeniya, Sri Lanka. In addition to developmental biol-

ogy, she is involved in morphological and molecular taxonomic studies.

Amphib. Reptile Conserv. | http://redlist-ARC.org 089

March 2012 I Volume 5 | Number 2 | e41

Commons Attribution License, which pennits unrestricted use, distribution, and reproduction in any medium,

provided the original author and source are credited.

Amphibian & Reptiie Conservation 5(2):90-100.

Calotes nigrilabris Peters, 1860 (Reptilia: Agamidae:

Draconinae): a threatened highland agamid lizard in Sri Lanka

^A. A. THASUN AMARASINGHE, ^FRANZ TIEDEMANN, AND ^D. M. S. SURANJAN KARUNARATHNA

^Komunitas Konservasi Alam Tanah Timur, Jl. Kuricang 18 Gd.9No.47, Ciputat 15412, Tangerang, INDONESIA ^Naturhistorisches Museum Wien,

Herpetologische Sammlung, Burgring 7, A- 1010 Vienna, AUSTRIA ^Nature Exploration & Education Team, B-l/G-6, De Soysapura Elats, Mora-

tuwa 10400, SRI LANKA

Abstract. — Caiotes nigriiabris Peters, 1860 is an endemic arboreal agamid lizard species that is

found only in montane and submontane cloud forests above 1,400 m elevation in central highlands

of Sri Lanka. Here we redescribe this species based on the holotype, newly collected material, and

published literature. Observations on the ecology, natural history, reproduction, and behavior of C.

nigriiabris are noted. Two specimens of C. nigriiabris were recorded from Thangappuwa (-1000 m

a.s.l.) in the Knuckles massif in 2003 and may represent a differentiated population needing further

study. Current habitat destruction and pesticide use in local farming practices are suggested as

primary threats to this species. A key to identifying members of the genus Caiotes in Sri Lanka is

provided.

Key words. Behavior, Caiotes nigriiabris, conservation, ecology, natural history, sauria, taxonomy

Citation: Amarasinghe AAT, Tiedemann F, Karunarathna DMSS. 2011. Caiotes nigriiabris Peters, 1 860 (Reptilia: Agamidae: Draconinae): a threatened

highland agamid lizard in Sri Lanka. Amphibian & Reptile Conservation 5(2):90-100 (e32).

Introduction

There are eighteen species of agamid lizards in Sri Lanka,

fifteen of which (83.3%) are endemic to the island (So-

maweera and Somaweera 2009; Manamendra-Arachchi

et al. 2006). Seven of these species belong to the genus

Caiotes, and five of which are endemic (C. ceylonensis

Muller, 1887; C. liocephalus Gunther, 1872; C. liolepis

Boulenger, 1885; C. nigriiabris Peters, 1860; C. desil-

vai Bahir and Maduwage, 2005) (De Silva 2006). The

remaining two Caiotes (C. caiotes Linnaeus, 1758 and

C. versicolor Daudin, 1802) are probably widespread

throughout Southeast Asia. According to published lit-

erature, the endemic Caiotes nigriiabris is a largely ar-

boreal species found only in montane and submontane

cloud forests above 1,400 m elevation (Das and De Silva

2005; Manamendra-Arachchi and Liyanage 1994). Its

conservation status is rare and vulnerable (Manamen-

dra-Arachchi and Liyanage 1994; lUCNSL and MENR

2007). However, Deraniyagala (1953) had reported a

specimen from Peradeniya (-650 m a.s.l), at a much

lower elevation than other known localities. Here we re-

describe this poorly known species based on the holotype

and newly collected specimens to provide more detailed

taxonomic information and proper identification of spe-

cies in this genus. This information is compiled into a

diagnostic key for the Sri Lankan members of the genus

Caiotes. Little ecological information is available for this

Correspondence. Email: Hhasun.taprobanica@ gmail.com

species, and further studies of its behavior and ecology

may be important for its conservation.

Methods and materials

The material examined is deposited at the NHMW,

Naturhistorisches Museum of Vienna, Vienna, Austria

and Wildlife Heritage Trust of Sri Lanka (WHT), Co-

lombo, Sri Lanka. Diagnoses and descriptions are based

on external morphology. The locality records for each

specimen include WHT specimen data, published local-

ity records as well as our observations during the past

decade (Fig. 1). All photographs and line drawings are

displayed with the photographer and artist initials: A.

Schumacher (AS), Thasun Amarasinghe (TA), Majintha

Madawala (MM), Gayan Pradeep (GP), and Vimukthi

Weeratunge (VW).

All measurements were taken to the nearest 0.1 mm

with dial calipers (Table 1). Scale counts: SUP, supral-

abials were counted from the first scale anterior to that

at angle of gape, not including the median scale (when

present); INF, infralabials were counted from first scale

posterior to mental, to angle of gape; DS, dorsal spines

were counted from first spine to last of mid-dorsal row;

CR, canthus rostralis were counted scales from rostral

scale along scale row passing over nostril to posterior

amphibian-reptile-conservation.org

090

November 2011 I Volume 5 I Number 2 I e32

Amarasinghe et al.

Figure 1. Current distribution patterns of C. nigrilabris (cen-

tral highland of Sri Lanka) (red circle: type locality and black

circle; other sightings).

end of supraciliary ridge; MDS, mid-dorsal scales were

counted from scale behind rostral to posterior margin of

the thigh; MBS, mid-body scales were counted from cen-

ter of mid-dorsal row forwards and downwards across

ventrals (this count is, however, made unreliable by

the unequal size and uneven arrangement of the lateral

scales); MVS, mid-ventral scales were counted from first

scale posterior to mental, to last scale anterior to vent;

SAT, spines around tympanum were counted from first

spine to last above tympanum. External measurements:

SVL, snout- vent length (distance between tip of snout to

anterior margin of vent); HE, head length (distance be-

tween posterior edge of mandible and tip of snout); HW,

head width (maximum width of head); DHE, dorsal head

length (distance between posterior edge of cephalic bone

and tip of snout); NEE, nostril-front eye length (distance

between anterior most point of orbit and middle of nos-

tril); UAE, upper-arm length (distance between axilla and

angle of elbow); EAE, lower-arm length (distance from

elbow to wrist with both upper arm and palm fiexed);

FE, finger length (distance between tip of claw and the

nearest fork); FEE, femur length (distance between groin

and knee); TBE, tibia length (distance between knee and

heel, with both tibia and tarsus fiexed); TE, toe length

(distance between tip of claw and nearest fork); AG, ax-

illa-groin length (distance between axilla and groin); SA,

snout-axilla length (distance between tip of snout and

axilla); TAE, tail length (measured from anterior margin

of vent to tail tip); PAE, palm length (taken from poste-

rior most margin of palm and tip of longest finger); FOE,

foot length (distance between heel and tip of longest toe,

with both foot and tibia fiexed); TBW, width of tail base

(greatest distance across the tail base); lOW, inter orbital

width (least distance between the upper margins of or-

bits); ED, eye diameter (horizontal diameter of orbit);

SEE, snout-front eye length (distance between anterior

most point of orbit and tip of snout); SBE, snout-back

eye length (distance between posterior most point of or-

bit and tip of snout); SET, snout-front tympanum length

(distance between anterior most point of tympanum and

tip of snout); TD, tympanum diameter (least distance be-

tween the inner margins of tympanum).

Calotes nigrilabris Peters, 1 860

Peters, W. C. H., Monatsberichte der Kdniglichen Akad-

emie der Wissenschaften zu Berlin, 1860: 183.

English Name: Ceylon black-cheek lizard or Dark-

lipped lizard; Sinhala Name: Kalii-kopul Katiissa or

Kalii-dekupul Katiissa.

Holotype: Male (99.8 mm SVE); Cat. no. NHMW

23355; Eoc. Newera Ellia: Ceylon (=Nuwara Eliya: Sri

Eanka); Coll. Unknown; Date. Unknown (see Amaras-

inghe et al. 2009 and Tiedemann et al. 1994; Fig. 2).

Other materials examined: WHT 0380A, WHT

0380B, WHT 0380C, WHT 0380D, Nagrak Division,

Nonpareil Estate, Horton Plains (06°46’ N 80°47’ E,

2135 m); WHT 0379, Kuda Oya, Eabugolla (07°0U N

80°44’ E, 1670 m); WHT 1555, WHT 2262, WHT 0536,

Hakgala (06°55’ N 80°49’ E, 1830 m).

Diagnosis

A row of 4, 5, or 6 laterally compressed spines above

the tympanum; lateral scales on the body directed back-

wards and downwards; dorsal and lateral scales on the

body much smaller than the ventral scales on the chest

and abdomen.

Figure 2. Dorsolateral view: holotype C. nigrilabris (Male)

(NHMW 23355) Nuwara Eliya, Sri Lanka (AS).

amphibian-reptile-conservation.org

091

November 2011 I Volume 5 I Number 2 I e32

Threatened highland agamid from Sri Lanka

Key to Sri Lankan species of genus Calotes

1 . No spines above the tympanum and lateral scales on the body pointing backwards and downwards

Calotes liocephalus

Spines above the tympanum present 2

2. Dorsal crest absent or less developed Calotes ceylonensis

Dorsal crest present and well developed 3

3. A row of laterally compressed spines above tympanum 4

Two separated spines above tympanum 5

4. Ventral scales larger than dorsal scales and scales on sides pointing backwards and downwards

Calotes nigrilabris

Ventral scales not larger than dorsal scales and scales on sides pointing backwards and upwards

Calotes calotes

5. Scales on sides pointing backwards and upwards Calotes versicolor

Scales on sides pointing backwards and downward 6

6. Gular sac present with black bands Calotes desilvai

Gular sac present without black bands Calotes liolepis

Description

(Based on the holotype and WHT collection). Length of

head one and half times its width; snout slightly longer

than orbit; rostral small, nasal rather large, forehead con-

cave; cheeks swollen in the adult male; upper head scales

unequal, smooth; 8 to 10 scales in canthal row, canthus

rostralis and supraciliary edge sharp; a row (3-6 spines)

of laterally compressed spines starting from above the

tympanum and extending posteriorly beyond it; diameter

of tympanum about half that of the orbit. Supralabials,

9-11; infralabials, VIII-IX (Fig. 3). Body laterally com-

pressed; dorsal scales more or less distinctly keeled,

pointing backwards and downwards (Fig. 4), except the

upper two or three rows with scales smaller than the ven-

trals, pointing directly backwards, strongly keeled, and

mucronate. Gular sac not developed, gular scales keeled,

as large as the ventrals; a short oblique pit or fold in front

of the shoulder covered with small granular scales. Nu-

chal and dorsal crests continuous, moderately developed,

composed of 17-27 lanceolate spines gradually diminish-

ing in size; the longest spines on the neck do not equal

the diameter of the orbit; female with a lower crest and a

mere ridge posteriorly. Limbs moderate; third and fourth

fingers equal or fourth finger a little longer than the third.

Relative length of fingers: 1<5<2<4<3 or 1<5<2<3<4.

Forth toe distinctly longer than the third. Relative length

of toes: 1<2<5<3<4. The hind limb reaches to the orbit

or the temple. Tail long and slender; in the adult male it

is markedly swollen at the base, with large, thick, keeled

scales.

Color pattern

(Based on our observations of live specimen; not collect-

ed). The body color is green with whitish, black-edged,

transverse bars or spots. Head marked with black; upper

lips and cheeks usually with a black streak or separated

from the eye by a white streak or with a pale bluish-green

stripe running from ear to shoulder; underside of the head

greenish- white, sometimes reddish-brown vertebral band

present or absent; base of the tail dark olive or brown

with darker-bordered light band or spots (Fig. 5).

Distribution and habitat

Calotes nigrilabris is endemic to Sri Lanka and had only

been recorded from montane and submontane cloud for-

ests above 1,400 m elevation in the central highlands.

However, examination of additional specimens reveals

that Calotes nigrilabris also occurs in the Horton Plains

(Kirigalpotta, -2200 m), which are grasslands around

Nuwara Eliya, Hakgala. Thus, C. nigrilabris is the only

Calotes species to occur in tropical high altitude open

grasslands (Bahir and Surasinghe 2005). According to

our observations C. nigrilabris is recorded from: Hor-

ton Plains National Park (06°46’ N 80°47’ E, ele. 2130

m); Kuda Oya, Eabugolla (07°0L N 80°44’ E, ele. 1670

m); Hakgala (06°55’ N 80°49’ E, ele. 1830 m); Nuwara

Eliya (06°57’ N 80°47’ E, ele. 1710 m); Piduruthalagala

(06°59’ N 80°46’ E, ele. 2300 m); Eabukele (07°0L N

80°42’ E, ele. 1525 m); Pattipola (06°5L N 80°50’ E,

amphibian-reptile-conservation.org

092

November 2011 I Volume 5 I Number 2 I e32

Amarasinghe et al.

Figure 3. Lateral side view (head scalation): male C. nigri-

labris (WHT 2262) Hakgala, Sri Lanka (Scale bar = 10 mm)

(TA).

Figure 4. Mid body lateral scales pointing backwards and

downwards of the male C. nigrilabris (WHT 0379) Labugolla,

Sri Lanka (Scale bar = 1 mm) (TA).

ele. 1890 m); Ohiya (06°49’ N 80°50’ E, ele. 1800 m);

Kandapola (06°59’ N 80°50’ E, ele. 1920 m); and Ragala

(06°59’ N 80°47’ E, ele. 1980 m).

Although the Dumbara population of Calotes nigri-

labris has long been recognized (Deraniyagala 1953), it

has not been compared critically with the populations

of the Central Hills. Unfortunately, the specimens from

Gammaduwa in the Dumbara Hills, deposited by De-

raniyagala in the National Museum of Sri Eanka, Co-

lombo, have since been lost. However, Erdelen (1984)

mentioned that he had no evidence of this species from

the Knuckles, in contrast to Deraniyagala (1953). Nev-

ertheless, we located C. nigrilabris from Thangappuwa

(-1000 m a.s.l.) in the Knuckles Region in 2003 and ob-

served two individuals (SVE 139.4 mm and 140.1 mm).

In ongoing research, we are working to clarify whether

these two populations are separate species.

Hemipenis morphology

There has been no serious attempt to classify agamid

lizards based on the morphological characters of the

hemipenis, even though there is an enormous diversity in

hemipenal morphology. The hemipenis of C. nigrilabris

seems less differentiated as compared to C. ceylonensis

(Karunarathna et al. 2009) and C. liocephalus (Amaras-

inghe et al. 2009). The hemipenis of Calotes nigrilabris

is well developed. The pedicel is slightly shorter than the

head; below the head, it is broadened out into two shal-

lowly concaved shoulders; there are no spines. The head

is quadrangular in shape. It is shallowly divided longitu-

dinally into four lobes, two being slightly larger than the

others. The surface of the head is pitted in a reticulating

pattern, the pits being larger on the outside and diminish-

ing in size towards the divisions between the lobes (Fig.

6 ).

Reproduction

The female digs a nest hole in the ground and deposits

two eggs in December (Deraniyagala 1953) and Taylor

(1953) observed two ova in each oviduct and the eggs

were 23 mm x 13 mm in size. We observed oviposition at

Horton Plains National Park in March 2010. The female

laid three eggs in the nest hole; sizes of the eggs were

17.5 mm x 10.1 mm, 17.8 mm x 10.8 mm, and 19.5 mm

X 10.2 mm (average size: 18.3 mm x 10.4 mm). In Sep-

tember 2001, we observed another female ovipositioning

at Nuwara Eliya. That female also laid three eggs; sizes

of the eggs were 17.4 mm x 9.8 mm, 17.0 mm x 9.7 mm,

and 17.1 mm x 9.7 mm (average size: 17.2 mm x 9.7

mm). A recent paper by Karunarathna et al. (2011) states

that female C. nigrilabris deposit 2-4 eggs at a time.

We have successfully hatched eggs in captivity.

Eggs were buried under soil in a screen-topped glass en-

closure. The above four eggs were half-buried in soil and

covered with leaf litter. The length of the enclosure mea-

sured 300 mm, width 150 mm, and height 100 mm. The

container holding the eggs was placed in a dark and cool

place (temperatures approximately 27.2-28. 5°C day time

and 25.7-26.4°C night). The relative humidity ranged

from 62%-78% during incubation. The surface soil was

generally kept dry, but occasionally about 50 ml of tap

water was sprayed in the hatching enclosure to maintain

a cool, humid environment similar to the original habitat.

November 2011 I Volume 5 I Number 2 I e32

amphibian-reptile-conservation.org

093

Threatened highland agamid from Sri Lanka

Table 1. Measurements (mm) and counts of the male holotype (NHMW 23355), three additional males, and five females of

Calotes nigrilabris (see measured material for specimen data).

Males (n=4)

NHMW 23355

WHT 0380C

WHT 1555

WHT 2262

Range

Mean ± SD

SVL

99.8

87.9

84.3

91.8

84 . 3 - 99.8

90.9 ± 5.8

34.1

34.0

31.9

33.6

31 . 9 - 34.1

33.4 ± 0.9

20.4

23.0

22.1

22.7

20 . 4 - 23.0

22.0 ± 1.0

DHL

25.7

25.0

26.0

24.4

24 . 4 - 26.0

25.3 ± 0.6

NFE

6.8

7.8

9.8

6.0

6 . 0 - 9. 8

7 . 6 ± 1.4

UAL

19.1

25.8

21.7

25.5

19 . 1 - 25.8

23.0 ± 2.8

LAL

22.0

17.7

17.2

19.9

17 . 2 - 22.0

19.2 ± 1.9

FLI

5.5

5.3

4.4

7.1

4 . 4 - 7. 1

5.6 ± 1.0

FL II

9.1

10.0

8.6

11.6

8 . 6 - 11.6

9 . 8 ± 1.1

FL III

14.7

15.1

10.2

15.8

10 . 2 - 15.8

13.9 ± 2.2

FL IV

14.4

13.9

12.4

15.9

12 . 4 - 15.9

14.1 ± 1.3

FL V

8.1

9.1

7.1

9.4

7 . 1 - 9.4

8.4 ± 0.9

FEL

23.2

28.7

23.3

29.6

23 . 2 - 29.6

26.2 ± 3.0

TBL

25.4

23.5

20.5

23.8

20 . 5 - 25.4

23.3 ± 1.8

TLI

6.5

10.5

5.6

8.2

5 . 6 - 10.5

7 . 7 ± 1.9

TLII

10.6

10.9

8.0

13.2

8 . 0 - 13.2

10 . 7 ± 1.8

TL III

17.0

12.1

15.0

18.5

12 . 1 - 18.5

15.6 ± 2.4

TL IV

20.8

14.4

16.7

21.7

14 . 4 - 21.7

18 . 4 ± 3.0

TL V

14.5

13.5

10.8

15.5

10 . 8 - 15.5

13 . 6 ± 1.8

46.8

42.5

39.3

44.7

39 . 3 - 46.8

43.3 ± 2.8

43.8

43.5

36.8

41.7

36 . 8 - 43.8

41.4 ± 2.8

TAL

285.7

225.0

broken

283

225 . 0 - 285.7

264.6 ± 28.0

PAL

22.1

23.2

15.1

19.9

15 . 1 - 23.2

20.1 ± 3.1

FOL

35.8

33.1

21.3

34.2

21 . 3 - 35.8

31.1 ± 5.7

TBW

10.5

11.1

10.7

12.0

10 . 5 - 12.0

11.1 ± 0.6

low

4.6

4.0

4.8

4.4

4 . 0 - 4.8

4.4 ± 0.3

8.4

7.0

8.6

8.3

7 . 0 - 8.6

8.1 ± 0.6

SFE

11.6

8.0

10.8

10.1

8 . 0 - 11.6

10.1 ± 1.3

SBE

18.6

18.0

19.7

18.9

18 . 0 - 19.7

18.8 ± 0.6

SFT

25.4

24.8

24.6

24.2

24 . 2 - 25.4

24.7 ± 0.4

3.5

6.0

3.7

5.4

3 . 5 - 6.0

4 . 6 ± 1.1

SUP

9-10

9.5 ± 0.5

INF

8-10

9.0 ± 0.7

MDS

59-66

63.3 ± 2.7

9-10

9.8 ± 0.4

MBS

48-58

51.3 ± 4.0

MVS

103

53-103

67.3 ± 20.7

amphibian-reptile-conservation.org

094

November 2011 I Volume 5 I Number 2 I e32

Amarasinghe et al.

Table 1 continued.

Females (n=5)

WHT 0380A

WHT 0380B

WHT 0380D

WHT 0379

WHT 0536

Range

Mean ± SD

SVL

71.0

71.8

74.9

77.3

70.7

70 . 7 - 77.3

73.1 ± 2.6

24.1

24.4

24.8

23.9

22.6

22 . 6 - 24.8

24.0 ± 0.7

14.1

13.5

13.6

13.4

13 . 4 - 14.1

13.7 ± 0.3

DHL

19.1

18.5

19.5

19.4

18.3

18 . 3 - 19.5

19.0 ± 0.5

NFE

5.2

6.4

5.8

4.7

4 . 7 - 6.4

5.5 ± 0.6

UAL

19.8

19.1

20.3

21.8

20.6

19 . 1 - 21.8

20.3 ± 0.9

LAL

17.3

14.7

17.2

15.8

15.3

15 . 3 - 17.3

16.1 ± 1.0

FLI

6.5

6.3

5.4

6.9

6.0

5 . 4 - 6.9

6.2 ± 0.5

FLII

10.0

10.6

7.6

9.4

10.1

7 . 6 - 10.6

9.5 ± 1.0

FLIII

12.6

14.3

10.1

12.7

12.6

10 . 1 - 14.3

12 . 5 ± 1.3

FL IV

11.9

13.9

10.8

11.8

11.6

10 . 8 - 11.9

12 ± 1.0

FL V

8.8

8.5

7.8

7.9

8.0

7 . 8 - 8. 8

8.2 ± 0.4

FEL

25.1

24.7

23.6

25.6

22.7

22 . 7 - 25.6

24.3 ± 1.1

TBL

19.3

18.7

18.8

20.1

18.9

18 . 7 - 20.1

19.2 ± 0.5

TLI

5.8

6.8

4.6

6.0

5.5

4 . 6 - 6.8

5.7 ± 0.7

TLII

8.5

10.8

7.7

8.8

8.1

7 . 7 - 10.8

8 . 8 ± 1.1

TL III

14.2

14.7

12.9

16.0

13.2

12 . 9 - 16.0

14 . 2 ± 1.1

TL IV

17.7

19.1

21.7

18.1

16.1

16 . 1 - 21.7

18.5 ± 1.9

TLV

12.4

13.1

9.6

12.0

9 . 6 - 13.1

11 . 8 ± 1.2

35.6

34.8

37.1

38.6

35.9

34 . 8 - 38.6

36.4 ± 1.3

34.8

33.7

33.2

33.7

29.6

29 . 6 - 34.8

33.0 ± 1.8

TAL

205

225

270

247

225

205-270

234.4 ± 22.2

PAL

16.7

15.6

13.8

15.6

18.7

13 . 8 - 18.7

16.1 ± 1.6

FOL

26.6

30.7

20.7

28.0

26.6

20 . 7 - 30.7

26.5 ± 3.3

TBW

6.8

9.1

7.4

8.4

6.8

6 . 8 - 9. 1

7.7 ± 0.9

low

3.5

3.2

2.4

3.4

3.8

2 . 4 - 3. 8

3.3 ± 0.5

6.3

6.5

7.5

6.5

7.5

63 - 1.5

6.9 ± 0.5

SFE

8.8

7.8

9.9

8.8

7 . 8 - 9.9

8.8 ± 0.7

SBE

15.4

15.1

16.8

15.4

15.6

15 . 1 - 16.8

15.7 ± 0.6

SFT

19.2

19.0

19.8

18.7

18.4

18 . 4 - 19.8

19.0 ± 0.5

3.8

3.7

3.4

3.6

3 . 4 - 3. 8

3.6 ± 0.1

SUP

9-11

9.8 ± 0.7

INF

8-9

8.8 ± 0.4

MDS

59-71

48.8 ± 7.3

8-10

9.0 ± 0.9

MBS

47-53

50.4 ± 2.5

MVS

51-64

58.8 ± 4.5

17-24

20.8 ± 2.6

SAT

4-5

4.4 ± 0.5

amphibian-reptile-conservation.org

095

November 2011 I Volume 5 I Number 2 I e32

Threatened highland agamid from Sri Lanka

Figure 5. Mature male C. nigrilabris (Nuwara Eliya) (black patch shown in cheek and small gular sac) (VW).

The lid of the container was close-fitting to deter preda-

tors (ants, etc.) and occasionally opened to spray water.

The juveniles emerged after 69 days. The emerging

hatchings waited approximately one hour, with snouts

extended from their shells, before rapidly exiting the egg.

The newly emerged juveniles ranged from 48.1-53.6 mm

in SVL and 2. 5-3. 2 g in weight (Table 2). After emerging

from their eggs, they were very active, running in circles

around the tank 10-15 times. We regularly provided small

earthworms, juvenile cockroaches, and termites. During

their first two days, these hatchlings only fed on earth-

worms and ate after breaking the prey into small parts.

On the third day, these animals refused earthworms and

only feed on juvenile cockroaches. They never fed on ter-

mites. Each individual ate 5-8 juvenile cockroaches per

day. After approximately 10 days, the hatchlings were re-

leased in good condition to the original habitat.

Table 2. Measurements (mm) and weight (WT) in grams of

hatchling Calotes nigrilabris in captivity (CH: Character).

(1)

(2)

(3)

(4)

Range

Mean ± SD

SVL

49.7

48.1

51.3

53.6

48.1-53.6

50.7 ±2.0

11.2

12.1

11.6

11.9

11.2-12.1

11.7±0.3

28.6

29.6

28.9

30.1

28.6-30.1

29.3 ± 0.6

2.6

2.8

2.5

3.2

2.5-3.2

2.8 ±0.3

Behavior

Fernando (1998) mentioned that male C. nigrilabris

gave a short hiss when handled. We also noted this hiss-

ing several times while handling this species. It is a very

Figure 6. Left hemipenis (lateral aspect) in C. nigrilabris

(WHT 1555) (TA).

amphibian-reptile-conservation.org

096

November 2011 I Volume 5 I Number 2 I e32

Amarasinghe et al.

Figure 7. Female C. nigrilabris on a Rhododendron arboretum bush (Horton Plains NP) (GP).

short, unrepeated “chik” sound, and it was only produced

by males.

Hatchlings are mostly found on bushes of Cymbo-

pogon sp., Panicum sp., Ulex europaeus, and Strobilan-

thes sp. and are typically light green. When disturbed or

danger approaches, these hatchlings take cover in an ad-

jacent bush. Mature individuals typically lie on endemic

Rhododendron arboreum shrubs and when disturbed, or

danger approaches, quickly jump into a nearby Cymbo-

pogon sp., Panicum sp., Strobilanthes sp., or Ulex euro-

paeus for refuge. This agamid lizard is usually sub-arbo-

real and inhabits tree trunks, hedges, and shrubs (Fig. 7)

where it hunts insects and earthworms by day (Das and

De Silva 2005).

Males are highly territorial and we observed terri-

torial fighting many times on tree trunks (Horton Plains

NP, Seetha Eliya, Pattipola, Agarapatana, Nuwara Eliya,

Eabukele, Haggala, and Ramboda). We never observed

the appalling, struggling, and chasing stages of combat

described by Karunarathna and Amarasinghe (2008).

During the “savaging stage,” they bite both fore and hind

limbs, cheeks, and nuchal crest of each other. They never

chased each other around the trunk while “savaging.”

Most often, they fight in open areas and the defeated indi-

vidual jumps down from the tree and escapes.

Conservation status

According to Erdelen (1988), the average population

density of C. nigrilabris was 220 individuals per hect-

are in Nuwara Eliya, and the population sizes and per-

centages of males, females, and juveniles were mostly

stable in Nuwara Eliya. According to Karunarathna et al.

(2011), the populations of C. nigrilabris are declining.

The official conservation status of the species is Vulner-

able (lUCNSE and MENR 2007).

Discussion

The threats to C. nigrilabris appear to stem largely from

habitat fragmentation. The impact of fragmentation

could be exacerbated by the fact that many important

amphibian-reptile-conservation.org

097

November 2011 I Volume 5 I Number 2 I e32

Threatened highland agamid from Sri Lanka

Figure 8. Typical forest and shrub habitat of C. nigrilabris (Horton Plains NP) (GP).

montane forest fragments are surrounded by agricultural

plantations (Fig. 8). Additionally, vegetable cultivation

in Sri Lanka involves the intensive and indiscriminate

application of pesticides (Erdelen 1984; Bahir and Sur-

asinghe 2005). These fast-moving lizards are susceptible

to mortality on roads (Fig. 9), and many hydropower

projects and rapid urbanization are continuing to modify

and fragment forest habitats. Additionally, C. nigrilabris

has a number of predators, including the Sri Lanka whis-

tling thrush {Myophonus blighi). Jungle crows (Corvus

macro rhynchos). Greater coucal (Centropus sinensis),

and feral cats {Felis catus), which were all recorded in

our study areas (Karunarathna and Amarasinghe 2008;

De Silva 2006; Warakagoda 1997). The crows are prob-

lematic because the local visitors to Horton Plains Na-

tional Park leave their garbage, which has encouraged

the migration and permanent settlement of Jungle crows

in Horton Plains NP. Therefore, these crows are a threat

for endemic C. nigrilabris, as well as other local reptiles.

The ecological and behavioral status of C. nigrila-

bris has been previously investigated by Erdelen (1978,

1984, 1988), who focused on population dynamics and

distribution of the genus Calotes in Sri Eanka, and by

Manamendra-Arachchi and Eiyanage (1994), who dis-

cussed the zoogeography of the Sri Eankan agamids.

The complete ovipositional behaviors of Calotes calotes

(Gabadage et al. 2009), Calotes versicolor (Amarasinghe

and Karunarathna 2007), Calotes nigrilabris (Karunara-

thna et al. 2011), Calotes liocephalus (Amarasinghe and

Karunarathna 2008), Calotes ceylonensis (Pradeep and

Amarasinghe 2009), and Calotes liolepis (Karunarathna

et al. 2009) are documented. However, ovipositional data

is lacking for C. desilvai.

According to Manamendra-Arachchi et al. (2006),

the lowlands (elevation -500 m) of the Mahaweli River,

which separates the Dumbara Hills (= Knuckles Hills)

from the Central Mountains, appears to have served as

a barrier to the dispersal of highland species between

the two mountain ranges. Therefore, genetic surveys of

these morphologically-defined populations are needed

to identify their evolutionary histories. If the Knuckles

Figure 9. Road killed sub-adult female C. nigrilabris (Horton

Plains NP) (MM).

amphibian-reptile-conservation.org

098

November 2011 I Volume 5 I Number 2 I e32

Amarasinghe et al.

population is a distinct species, then that species could

be critically endangered due to habitat fragmentation by

Cardamom (Elettaria cardamomum) and tea {Camel-

lia sinensis) cultivations, which also often involve the

intensive and indiscriminate application of pesticides.

Conservation breeding programs may be needed if the

population sizes of the species continue to decline in its

natural habitat.

Acknowledgments. — Wq would like to express our

sincere thanks to Rohan Pethiyagoda (WHT), Sudath

Nanayakkara (WHT), and Craig Hassapakis (ARC), who

helped in diverse ways to enrich this work. We thank

Toshan Peries, Than Abeywardene, Panduka Silva,

Anushka Kumarasinghe, Asanka Udayakumara, Niran-

jan Karunarathna, Chamila Soysa, Mahesh C. De Silva,

and members of the YZA for various help in the field.

Finally, we would like to thank Vimukthi Weeratunge

(lUCN Sri Lanka), Majintha Madawala (YZA), Gayan

Pradeep (YZA), and A. Schumacher (NHMW) for pro-

viding photographs.

Literature cited

AmarasingheAAT, Karunarathna DMSS. 2007. Beobachtun-

gen zum Eiablageverhalten der Indischen Schonechse Calo-

tes versicolor (Daudin, 1 802) (Reptilia; Agamidae) in einem

anthropogenen Biotop in Sri Lanka. Sauria 29(3):27-30.

AmarasingheAAT, Karunarathna DMS S . 2008 . Observations

on the ovipositional behavior of the Crest-Less Lizard Calo-

tes liocephalus (Reptilia; Agamidae) in the Knuckles For-

est Region of Sri Lanka. Asiatic Herpetological Research

11:13-16.

Amarasinghe AAT, Karunarathna DMSS, Gab adage DE.

2009. Current status of Calotes liocephalus Gunther, 1872

(Reptilia; Agamidae) of Sri Eanka. Journal of Threatened

Taxa l(ll);553-557.

Amarasinghe AAT, Manthey U, Stockli E, Ineich 10, Kul-

LANDER S , T lEDEMANN F, McC ARTHY C , GaB ADAGE DE . 2009 .

The original descriptions and figures of Sri Eankan Agamid

Eizards (Squamata: Agamidae) of the 18th and 19th centu-

ries. Taprobanica 1(1):2-15 + 4 plates.

Bahir, mm, Surasinghe TD. 2005. A conservation assessment

of the agamid lizards of Sri Eanka. The Raffles Bulletin of

Zoology (Supplement) 12:407-412.

Das 1, De Silva A. 2005. Snakes and Other Reptiles of Sri Lan-

ka. New Holland Publishers, United Kingdom. 144 p.

Deraniyagala PEP. 1953. A Colored Atlas of Some Vertebrates

from Ceylon. Volume 2, Tetrapod Reptilia. Ceylon Govern-

ment Press / Ceylon National Museums, Colombo. 101 p.

De Silva a. 2006. Current status of the reptiles of Sri Eanka. In

Fauna of Sri Lanka: status of taxonomy, research and con-

servation. Editor, CNB Bambaradeniya. lUCN Sri Eanka,

Colombo. 134-163.

Erdelen W. 1978. Distribution patterns of the genus Calotes

(Sauria; Agamidae) of Sri Eanka. Loris 14(6);350-353.

Erdelen W. 1984. The genus Calotes (Sauria, Agamidae) in

Sri Eanka: distribution patterns. Journal of Biogeography

11:515-525.

Erdelen W 1988. Population dynamics and dispersal in three

species of agamid lizards in Sri Eanka: Calotes versicolor,

C. calotes, C. nigrilabris. Journal of Herpetology 22{\):A2-

52.

Fernando M. 1998. Hissing of Peters, 1860.

Sri Lanka Naturalist 2(3):3 1 .

Gabadage de, Amarasinghe AAT, Bahir MM. 2009. Notes on

the ovipositional behaviour of Calotes calotes (Einnaeus,

1758) (Reptilia: Agamidae) in Sri Eanka. Herpetotropicos

5(l):21-24.

lUCNSE & MENR. 2007. The 2007 Red List of Threatened

Fauna and Flora of Sri Lanka. lUCN Sri Eanka, Colombo.

148 p.

Karunarathna DMSS, Amarasinghe AAT. 2008. An observa-

tion of the jungle crow (Aves; Corvidae) feeding on Ceylon

Pygmy Lizards, Cophotis ceylanica (Reptilia: Agamidae) at

Horton Plains NP in Sri Lanka. Sauria 30(4): 59-62.

Karunarathna DMSS, Amarasinghe AAT. 2009. Notes on

the territorial behavior of Otocryptis wiegnianni Wagler,

1830 (Reptilia: Agamidae: Draconinae). Herpetotropicos

4(2):79-83.

Karunarathna DMSS, AmarasingheAAT, Stockli E. 2009.

Taxonomical and biological study on Calotes ceylonensis

Muller, 1887 (Reptilia; Agamidae) of Sri Eanka. Bonner zo-

ologische Beitrdge 56(4):229-238.

Karunarathna DMSS, Bandara IN, Chanaka AWA. 2009.

The ovipositional behaviour of the endemic whistling liz-

ard Calotes liolepis Boulenger, 1885 (Reptilia: Agamidae)

in the Knuckles forest region of Sri Eanka. Acta Herpeto-

logica 4(l):47-56.

Karunarathna DMSS, Pradeep WAADG, Pe abotuwage PIK,

De Silva MC. 2011. First report on the ovipositional behav-

iour of Calotes nigrilabris Peters, 1860 (Reptilia: Sauria:

Agamidae) from the Central massif of Sri Eanka. Russian

Journal of Herpetology 1 8(2); 111-118.

Manamendra-ArachchiK, De Silva A, Amarasinghe T. 2006.

Description of a second species of Cophotis (Reptilia:

Agamidae) from the highlands of Sri Eanka. Lyriocephalus

6: (Supplement) 1:1-8.

Manamendra-Arachchi K, Eiyanage S. 1994. Conservation

and distributions of the agamid lizards of Sri Eanka with

illustrations of the extant species. Journal of South Asian

Natural History l(l):77-96.

Peters WCH. 1860. liber einige interessante Amphibien,

welche von dem durch seine zoologischen Schriften riih-

mlichst bekannten osterreichischen Naturforscher Profes-

sor Schmarda wahrend seiner auf mehrere Welttheile aus-

gedehnten, besonders auf wirbellose Thiere gerichteten,

naturwissenschaftlichen Reise, mit deren Veroflfentlic-

hung Hr. Schmarda gegenwartig in Berlin beschaftigt ist,

auf der Insel Ceylon gesammelt wurden. Monatsberichte

der Kdniglichen Akademie der Wissenschaften zu Berlin

amphibian-reptile-conservation.org

099

November 2011 I Volume 5 I Number 2 I e32

Threatened highland agamid from Sri Lanka

(April); 182-186.

Pradeep WAADG, Amarasinghe AAT. 2009. Ovipositional

behavior of Calotes ceylonensis Muller, 1887 (Reptilia;

Agamidae) observed at the central province of Sri Lanka.

Taprobanica l(l):24-27.

SoMAWEERA R, SoMAWEERA N. 2009. Lizards of Sri Lanka: a

colour guide with field keys. Edition Chimaira / Serpent’s

Tale NHBD, Germany. 304 p.

Taylor EH. 1953. A review of the lizards of Ceylon. The Uni-

versity of Kansas Science Bulletin 35(12): 1525-1585.

T lEDEMANN F, Haupl M, Grillitsch H. 1 994. Katalog der Typen

der Herpetologischen Sammlung nach dem Stand vom Jan-

ner 1994, Teil IE Reptilia. Kataloge der wissenschaftlichen

Sammlungen des Naturhistorischen Museums in Wien, Wien

(Naturhistorisches Museum Wien), 10 (Vertebratad). 110 p.

Warakagoda D. 1997. Some observations on the Sri Eanka

Whistling Thrush. OB C Bulletin 26:33-34.

Manuscript received: 31 October 2010

Accepted: 25 April 2011

Published: 20 November 2011

A. A. THASUN AMARASINGHE is a Sri Eankan herpetolo-

gist. He is the chairman of Komunitas Konservasi Alam Tanah

Timur - Indonesia, Editor-in-Chief of Taprobanica: The Jour-

nal of Asian Biodiversity, and a member of the Crocodile Spe-

cialist Group in lUCN/SSC. Currently he is involved in ecology

and taxonomy projects in Sri Eanka and Indonesia.

DR. FRANZ TIEDEMANN specializes in herpetology and

habitat mapping and is also president of the Austrian Society

of Herpetology (1990 to 2002), and member of the editorial

board of the journal Herpetozoa (2002). He is currently holds

the position of Honorary Staff member at the Naturhistorisches

Museum Wien, Austria (Natural History Museum of Austria),

and member of the Tortoise and Freshwater Turtle Specialist

Group lUCN/SSC in 1991-1992.

D. M. S. SURANJAN KARUNARATHNA is a field biologist

conducing research on amphibian and reptile ecology, and pro-

moting conservation awareness of the importance of biodiver-

sity among the Sri Eankan community. He began his career and

wildlife research in 2000, as a member of the Young Zoologists’

Association (YZA) based in the department of the National Zo-

ological Gardens of Sri Eanka. He worked as an ecologist for

the lUCN Sri Eanka County Office and is an active member of

many specialist groups in the lUCN/SSC.

amphibian-reptile-conservation.org

100

November 2011 I Volume 5 I Number 2 I e32

Attribution-NonCommercial-NoDerivs 3.0 Unported License, which permits unrestricted use for non-commercial

and education purposes only provided the original author and source are credited.

Amphibian & Reptiie Conservation 5(2):101-113.

Territorial and site fidelity behavior of Lyriocephalus scutatus

(Agamidae: Draconinae) in Sri Lanka

^Imesh Nuwan Bandara

^“Ellangaawa” Unity care for Community & Nature, No: 1/112, Hapugoda, Ambatenna, SRI LANKA 20136;

Youth Exploration Society of Sri Lanka, PO. box 82, Peradeniya, SRI LANKA

Abstract — ^This study on territorial behavior of Lyriocephaius scutatus suggests that territorial be-

havior is an important component of the life history of the species. Lyriocephaius scutatus belongs

to the monotypic genus Lyriocephaius, and apparently its uniqueness, placing it in its own genus,

extends to its strange behavior and atypical site fidelity. To understand this territorial behavior, two

populations were observed while continuously recording other factors influencing territorial and

site fidelity behaviors. Individual lizards performed various behaviors in their daily active periods on

tree trunks and on the ground. They also exhibited highly specific synchronized territorial behavior

among other individuals in the same population. Behavioral patterns differed between males and fe-

males, and the degree of “aerial horizontal distribution” of L. scutatus seems to be a novel behavior

among lizards. Individual L. scutatus are highly territorial over other individuals of the same sex, as

adult males observed in the study sites solely performed their territorial displays on a specific tree,

whereas females occupied the largest territories.

Key words. Territorial behavior, Lyriocephaius scutatus. Lyre head lizard, Sri Lanka

Citation: Bandara IN. 2012. Territorial and site fidelity behavior of Lyriocephaius scutatus (Agamidae: Draconinae) in Sri Lanka. Amphibian & Reptile

Conservation 5{2)-A0t-tt3 (e56).

Introduction

Sri Lanka is a continental island endowed with high her-

petofaunal diversity and endemism. Two-hundred and

seven species of reptiles have been described from Sri

Lanka and more than half are endemic to the island (So-

maweera and Somaweera 2009). The agamid lizard fau-

na of Sri Lanka is comprised of 18 species in six genera,

15 of which are endemic (Bahir and Surasinghe 2005;

Samarawickrama et al. 2006): Calotus (six species; four

endemic), Ceratophora (five species; all endemic), Co-

photis (two species; both endemic), Lyrocephalus (one

endemic species), Otocryptis (two species; both endem-

ic), and Sitana (one species of unclear taxonomic sta-

tus). Of these genera, Lyrocephalus, Ceratophora, and

Cophotis are are considered to be relict lineages because

they are confined to Sri Lanka.

In spite of the uniqueness of the lizard fauna of Sri

Lanka, little is known with regard to the behavior, ecol-

ogy, and natural history for most of the agamid species.

This is particularly true with regard to territoriality, even

though males of most species are presumed to be terri-

torial. Among the short observation notes on territorial

behavior of Sri Lankan agamids are works by Derani-

yagala (1931, 1953), Smith (1935), Bambaradeniya et

al. (1997), and Karunarathna and Amarasinghe (2008).

However, there have been no long-term studies on ter-

Correspondence. Email: imeshnul @ gmail.com

ritorial behavior of any Sri Lankan agamid lizard. One

species, Lyrocephalus scutatus, is of particular interest

because it is the only species of the genus and is endemic

to Sri Lanka (Figs. 1 and 2). Several authors, (Derani-

yagala 1931, 1953; de Silva et al. 2005; Manamendra-

Arachchi 1998) reported L. scutatus to have territorial

behaviors with males intimidating each other by opening

wide their blood-red mouths showing their long sharp

teeth and shaking their heads. Additionally, when threat-

ened, they would lie motionless on their sides feigning

death. A better understanding of these behaviors is nec-

essary to more completely appreciate the unique lizard

fauna of Sri Lanka and to aid in its conservation. Hence,

the present study examines territorial and site-fidelity be-

havior of the endemic lizard L. scutatus.

Methods and materials

Study area

The study took place in the Gannoruwa Forest Reserve

[GFR] (T 17’ N, 80° 36’ E) in Kandy district in the Cen-

tral Province of Sri Lanka (Fig. 3; modified from Wick-

ramasinghe 2006). The reserve is a remnant forest patch

covering an area of -250 acres and is surrounded by vil-

lages. The vegetation within the GFR can be grouped into

amphibian-reptile-conservation.org

101

October 2012 I Volume 5 I Number 2 I e56

Bandara

natural forest, naturalized plantations (i.e., abandoned

cocoa, tea, coffee, Artocarpus hetewphyllus, etc.), grass-

lands, and mahogany plantations. Home gardens com-

prise most of the anthropogenic ecosystems bordering

the reserve. Observations were made at two sites within

the GFR. Site A at Pallegama (07*^ 28’ N and 80*^ 60’ E)

is a high canopy home garden that is very well shaded

by the common tree Myristica fragrance (Fig. 4) with a

moderate to steep slope (30° average). Site B at Yatihala-

gala (07° 36’ N and 80° 52’ E) is also a shady habitat, but

with greater human interference than study Site A since it

is nearer to human settlements (Fig. 5). Garmin (GPS 12)

was used to obtain geographical coordinates and Brun-

ton clinometers (Brunton Company, USA) were used for

measuring slope.

Methods

Detailed studies started in mid October, 2005, and were

conducted until late February, 2006. Both field sites were

partitioned into a grid of 1 x 1 m quadrats using small

PVC stumps to mark the coordinates so locations of liz-

ards could be determined within 0.25 m. Two template

grid maps were created, one for each of the study sites.

Each lizard observed was captured, sexed, measured, and

given an identifying name. To permit identification of in-

dividual lizards from several meters away, all individu-

als observed and captured, within the study areas, were

temporarily marked using loose elastic bands of various

colors placed on the waist. Three reproductive classes

were recorded: adult males, adult females, and subadults.

Adult males and females were defined as individuals that

were sexually mature (i.e., >80 mm snout to vent length

[SVL] and with fully grown rostral knob and crest). Sub-

adults were defined as individuals that were not in breed-

ing condition (i.e., <80 mm SVL and less developed ros-

tral knob and crest). Direct visual observation of natural

populations was aided, when necessary, by the use of a

pair of Nikon 10 x 8 binoculars. Focal population sam-

pling was conducted by observing the entire population

continuously for 20 to 60 minutes; thus the observed fo-

cal time for individual animals of a particular population

was equal. If a particular animal was not located during

the entire sampling it was considered “Not Observed.”

In order to gather detailed information on spatial dis-

tribution, censuses were conducted three times a month

by traversing the entire field site. Trees in which lizards

were observed were scanned throughout the day (0600 to

1800 hr) and the locations of all lizards (marked or un-

marked) were recorded. All behaviors observed, includ-

ing both those exhibited in isolation and those directed

towards other individuals, were recorded and all individ-

uals involved in social interactions were noted. A total of

110 hours was spent performing the censuses.

Herein, an individual lizard’s territory is considered

to be the area that encompasses all positions of the liz-

ard, day and night. Thus, all locations where individu-

als were observed during the study period (including

incidentally observed individuals) were recorded and

mapped for the calculation of the size of the territory.

Territories are graphically displayed as polygons with

inside angles <180° hand-drawn around the outermost

observed coordinates. In addition, all woody surfaces of

trees where lizards were recorded were added to the area

of the territory using average cylindrical area represent-

ing the trunk of a tree (Philibosian 1975). Since we have

repeated measurements of the same individuals on dif-

ferent days, and multiple individuals from the same site

(spatial autocorrelation) data were analyzed statistically

as a linear mixed effects model using software R-2.9.0-

win32. Microsoft Office Excel 2007 was used for the

graphical display of data.

Results

Observed behaviors

A total of 180 focal animal samples were recorded from

12 marked individuals (six males, three females, and

three subadult males) on 15 days (including night visits)

over a six-month period in the pre-reproductive season

of these lizards. The marked population at Site A con-

sisted of five individuals (two males, one female, and two

subadults). The marked population Site B consisted of

seven individuals (four males, two females, and one sub-

adult). All behaviors demonstrated, including both those

exhibited in isolation and those directed towards other

individuals, are summarized in Table 1.

Table 1. Summary of commonly observed behaviors of lizards in their natural environment.

Behaviors Description

Body-lift

Gular Sac Display

Head-bob

Tail-wag

Still

Adjustment

Walking

Feeding

Uplift on all four limbs pushing body off surface followed immediately by deseent, repeated frequently; other lizards

may or may not be seen in the vieinity.

Gular sae is extended with lateral side eompression aeeompanying a Body-lift.

Relatively rapid up-and-down movement of the head or head and neek region only; gular sae may also be extended.

Undulating movement of tail.

Positioned on the surfaee without notable movements.

A simple ehange in still position.

Moving about in an area slowly.

Taking in a food item.

amphibian-reptile-conservation.org

102

October 2012 I Volume 5 I Number 2 I e56

Territorial behavior of Lyriocephalus scutatus

Figure 1. Lyrocephalus scutatus - male.

Figure 3. Map of Gannoruwa Forest Reserve - Kandy district

Sri Lanka.

Figure 2. A lizard threat pose - Body-lift, Gular Sac Display, and Head-bob.

amphibian-reptile-conservation.org

103

October 2012 I Volume 5 I Number 2 I e56

Bandara

Figure 4. Study site A - Gannoruwa Pallegama.

Figure 5. Study site B - Gannoruwa Yatihalagala.

amphibian-reptile-conservation.org

104

October 2012 I Volume 5 I Number 2 I e56

Territorial behavior of Lyriocephalus scutatus

Figure 6. Different behaviors observed: Body-lift, Gular sac display, and Head-bob with Body-lift.

Figure 7. Different behaviors observed: Tail wag. Adjustment, and Still.

amphibian-reptile-conservation.org

105

October 2012 I Volume 5 I Number 2 I e56

Bandara

Behaviors

Body lift

Cularsac dtsp^y

Mead bob

Tailwjg l«mc4

Adjustment

SUN

Walking

Feeding

06.(X>- 10.00 3.m. 10.00-14.00 14.00-18.00 P*n>. Time

Figure 8. Percentage of various behaviors displayed by L. scutatus according to time of day.

Figure 9. Percentage of time spent exhibiting various behaviors for the different reproductive groups of L. scutatus: males, females,

and subadults.

amphibian-reptile-conservation.org

106

October 2012 I Volume 5 I Number 2 I e56

Territorial behavior of Lyriocephalus scutatus

Cf aLOl

aL03

O aL02

Q aL05

aL04

Figure 10. Map of individual territories of lizards in site A.

Table 2. Percentage of overlap of territories between individuals according to reproductive category.