Published in the United States of America 2012 * VOLUME 5 * NUMBER 2 V/ , =-> V ■ •• - — • 2> -- < j-=r “17 m I- " : : ^ '■ lit' r-. ^ S r V ^ « '^1 * «U1 V :> a' ' V ■ ■ '"H. ; S N *'. SRI LAEfl^ ISSN: 1083-446X elSSN: 1525-9153 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 © Craig Hassapakis/Amphibian & Reptile Conservation Copyright: © 2011 Janzen and Bopage. This is an open-access article distributed under the terms of 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):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. 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 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 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. 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. 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 Copyright: © 201 1 Ariyasiri et al. This is an open-access article distributed under the terms of the Creative Com- 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 1 2 3 4 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 TL 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 BL -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. 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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 Copyright: © 2011 Bowatte and Meegaskumbura. This is an open- access article distributed under the terms of 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 = 2 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 Copyright: © 2012 Erdelen. This is an open-access article distributed under the terms of 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):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. 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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 Copyright: © 2012 Peabotuwage et al. This is an open-access article distributed under the terms of 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):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. 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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 Copyright: © 2012 Botejue and Wattavidanage. This is an open-access article distributed under the terms of 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):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 CF FE HG CU Ichthyophiidae Ichthyophis glutinosus ® X X X - Bufonidae Adenomus kelaartii ® X X X - Duttaphrynus melanostictus X X X X Microhyiidae Kaloula taprobanica X X X X Micwhyla rubra X X X - Ramanella variegata X X X - Dicrogiossidae Euphlyctis cyanophlyctis X X X X Euphlyctis hexadactylus X X X X Zakerana kirtisinghei ® X X X X Zakerana syhadrensis X X X X Hoplobatrachus crassus X X X X Nannophrys ceylonensis X X X - Nyctibatrachidae Lankanectes corrugatus ® X X X X Ranidae Hylarana aurantiaca X X X - Hylarana temporalis X X X - Rhacophoridae Pseudophilautus abundus ® X X - - Pseudophilautus cavirostris X X - - Pseudophilautus folicola X X - - Pseudophilautus hoipolloi ® X X X - Pseudophilautus popularis ® X X X X Pseudophilautus reticulatus ® X - - - Pseudophilautus stictomerus X X - - Polypedates cruciger ® X X X X Taruga longinasus X X - - 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 CF FE HG CU CF FE HG CU Pythonidae Uropeltidae Python molurus X X X X Rhinophis sp. X X - - Colubridae Viperidae Ahaetulla nasuta X X X - Daboia russelii X X X X Ahaetulla pulverulenta X - X - Hypnale hypnale X X X X Amphiesma stolatum X X X X Trimeresurus trigonocephalus ® X X - - Aspidura guentheri X X - - Agamidae Atretium schistosum X X - - Calotes calotes - X X X Balanophis ceylonensis X - - - Calotes liolepis X X X X Boiga ceylonensis X - X - Calotes versicolor - X X X Boiga forsteni X X X X Ceratophora aspera ™ X - - - Cercaspis carinatus X - X - Lyriocephalus scutatus X X - - Chrysopelea ornate X X X - Otocryptis wiegmanni X X X X Coelognathus helena X X X X Gekkonidae Dendrelaphis bifrenalis X - - - Cnemaspis silvula ® X X X - Dendrelaphis caudolineolatus X X - - Cnemaspis sp. X X - - Lycodon aulicus X - X X Geckoella triedrus X - - - Lycodon osmanhilli ® X X X X Gehyra mutilata - - X X Oligodon arnensis X X X X Hemidactylus depressus ® X X X - Oligodon sublineatus ® - X X X Hemidactylus frenatus X X X - Ptyas mucosa X X X X Hemidactylus parvimaculatus X X X - Sibynophis subpunctatus X X X X Lepidodactylus lugubris ™ X X X - Xenochrophis asperrimus ® X X - - Scincidae Xenochrophis piscator X X - - Eutropis carinata X X X X Cylindrophiidae Eutropis madaraszi X X - - Cylindrophis maculatus X X X - Lankascincus fallax ® X X X X Elapidae Lankascincus gansi X X - X Bungarus ceylonicus X X X - Lankascincus greeri ® X X X X Naja naja - X X X Varanidae Typhlopidae Varanus bengalensis - X X X Ramphotyphlops sp. X X - - Varanus salvator - X X X Typhlops sp. X X - - Bataguridae Melanochelys trijuga - X X X 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). 60 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 u c ro c 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. 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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. 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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 Copyright: © 2012 Samarawickrama et al. This is an open-access article distributed under the terms of 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):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 Copyright: © 2011 Amarasinghe et al. This is an open-access article distributed under the tenns of the Creative 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 HL 34.1 34.0 31.9 33.6 31 . 9 - 34.1 33.4 ± 0.9 HW 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 AG 46.8 42.5 39.3 44.7 39 . 3 - 46.8 43.3 ± 2.8 SA 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 ED 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 TD 3.5 6.0 3.7 5.4 3 . 5 - 6.0 4 . 6 ± 1.1 SUP 10 10 9 9 9-10 9.5 ± 0.5 INF 10 9 9 8 8-10 9.0 ± 0.7 MDS 65 59 66 63 59-66 63.3 ± 2.7 CR 10 10 10 9 9-10 9.8 ± 0.4 MBS 50 49 58 48 48-58 51.3 ± 4.0 MVS 57 56 103 53 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 HL 24.1 24.4 24.8 23.9 22.6 22 . 6 - 24.8 24.0 ± 0.7 HW 14.1 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 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 12.0 9 . 6 - 13.1 11 . 8 ± 1.2 AG 35.6 34.8 37.1 38.6 35.9 34 . 8 - 38.6 36.4 ± 1.3 SA 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 ED 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 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 TD 3.8 3.7 3.4 3.6 3.6 3 . 4 - 3. 8 3.6 ± 0.1 SUP 10 11 10 9 9 9-11 9.8 ± 0.7 INF 9 8 9 9 9 8-9 8.8 ± 0.4 MDS 59 64 62 59 71 59-71 48.8 ± 7.3 CR 9 10 8 8 10 8-10 9.0 ± 0.9 MBS 51 53 48 53 47 47-53 50.4 ± 2.5 MVS 61 64 57 61 51 51-64 58.8 ± 4.5 DS 24 23 21 19 17 17-24 20.8 ± 2.6 SAT 5 5 4 4 4 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). CH (1) (2) (3) (4) Range Mean ± SD SVL 49.7 48.1 51.3 53.6 48.1-53.6 50.7 ±2.0 HL 11.2 12.1 11.6 11.9 11.2-12.1 11.7±0.3 AG 28.6 29.6 28.9 30.1 28.6-30.1 29.3 ± 0.6 WT 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 Copyright: © 2012 Bandara.This is an open-access article distributed under the terms of the Creative Commons 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. Category Male Female Subadult Male <1.0% 40% 20% Female 48% 00% 11% Subadult 15% 18% 00% amphibian-reptile-conservation.org 107 October 2012 I Volume 5 I Number 2 I e56 Bandara Several different behaviors were observed during liz- ard activity periods (Figs. 6 and 7). Generally, display- ing would begin in the morning and continue for several hours until the displaying lizards would climb down from trees to the ground. In the evening, lizards would climb- up the trees and start displaying again until they would go to sleep at nightfall. When an individual lizard did not comedown from the tree, it remained there in the Still po- sition the entire day. When lizards were displaying they performed their behaviors in an upright position on the tree trunks. Body-lift, Gular Sac Display, Head-bob, and Tail-wag were frequently performed in the upright po- sition, however Head-bob, Body-lift, and Tail-wag were also performed on the ground while performing Walking or Feeding. The meeting of two different individuals was not ob- served during the six-month study period. On one occa- sion, a female was found with a male on the same tree, but no remarkable behaviors were observed between those two individuals, although the male did display its usual behaviors. Behavioral differences among reproductive groups Time spent performing the various behaviors differed with the time of the day (Fig. 8). Behaviors such as Head-bob, Body-lift, Gular Sac Display, and Still were common in the morning hours from 0600 to 1000 hr. Feeding was not observed during this morning time pe- riod and only a small amount of time was spent Walk- ing. Tail-wag and Adjustment were also performed in the morning. During daytime, from 1000 to 1400 hr. Walking increased to about 60% of the behavior of L. scutatus. All the other behaviors observed in higher proportion during k N 0 bL 06 CT bL ID Cf bL D8 cT bL07 bL09 Q bL 11 bL 12 Figure 11. Map of individual territories of lizards in site B. amphibian-reptile-conservation.org 108 October 2012 I Volume 5 I Number 2 I e56 Territorial behavior of Lyriocephalus scutatus the day gradually decreased in frequency as time passed. Evidently 90% of Feeding was done in the midday hours (1100-1230 hr). During the evening hours from 1400 to 1800 hr, Still was demonstrated by 70% of the individu- als observed, and all the other behaviors became rarer, especially Walking and Feeding. Overall, the behaviors exhibited by the lizards var- ied with time from morning to evening. Additionally, all individuals at a particular site would synchronize their behavior. For example, when a certain individual would begin the Gular Sac Display, all individuals at that par- ticular site would soon perform the Gular Sac Display Figure 12. Arboreal distributions of lizards in site A. as well. Normally the dominant male would initiate the display with other individuals following with the same A paired t-test showed that there is a significant difference in the patterns of behaviors between males and females (t = 3.10, p = 0.004). Not only were the behaviors shown by males and females markedly different, the percentages of time spent in each behavior differed as well (Fig. 9). Body-lift and Gular Sac Display were confined to males and Head-bob and Tail-wag were shared by both sexes, but males had a comparatively higher percentage. About 60% of the observed instances of Adjustment, Still, and Walking were performed by females. One instance of mating behavior of L. scutatus was observed in this study. The single observation was about 2.4 m above the ground at 0720 hrs in the morning on a Syzygium aromaticum tree with a girth of 42 cm. Copula- tion was maintained for three minutes, after which both individuals were observed in the same tree for the dura- tion of the day. Territoriality The size of territory differed among reproductive groups with females having the largest (264.94 + 59.8 m^), fol- lowed by males (178.72 + 32.1 m^), and then subadults (174.73 + 32.3 m^), although males and subadults had roughly equal sized territories (Figs. 10 and 11). A Paired Mest showed a significant difference between male and female territories (t = 2.38, p = 0.02). Territory size was not linked to the body size of the owner (t = 2.S, p = 0.008). On five occasions male territories were overlapped by approximately 40% by a female territory and on four Figure 13. Arboreal distributions of lizards in site B. amphibian-reptile-conservation.org 109 October 2012 I Volume 5 I Number 2 I e56 Bandara occasions male territories were overlapped by approxi- mately 20% by a subadult territory (Table 2). Only on one occasion did a male territory overlap another male territory, although this overlap involved less than 1.0% of each territory. On six occasions female territories were overlapped by approximately 48% by a male territory, and on a single occasion a female territory was over- lapped by approximately 11% by the territory of a sub- adult. On three occasions, subadult territories were over- lapped by approximately 15% by a male territory and on a single occasion a subadult territory was overlapped by approximately 18% by the territory of a female. Over- lap of territories among the same reproductive group was not observed in this study except on the single occasion of the two males with territories overlapping less than 1.0%. In fact, all males observed in the two study sites were on a tree with no other lizards present (marked with male symbol in Figs. 10 and 11), and they remained on “their” tree throughout the study period with the single exception of the lizard “bL 08” which was recorded oc- cupying two different trees. Males displayed only when they were on their particular tree. Females and subadults were recorded on several trees within their particular home range. Arboreal distribution As a group, the lizards of this study showed a previously unreported behavior of maintaining a particular level of height on the trees, especially while displaying. When the observed lizards climbed-up trees they all appear to stop at a similar and consistent elevation. In Site B all the individuals maintained an arboreal height of 2.5 m to 4.1 m, and since the area is rather flat their distribu- tion approximately paralleled the ground. It was only at these positions in the trees that the lizards performed syn- chronized display behaviors (Figs. 12 and 13). In Site A, which has a slope of 30°, the level of the height of lizards forms about a 60° angle to the ground. Interestingly, at Site A, when the dominant male started to adjust its posi- tion all other lizards at the site adjusted their positions, thus maintaining the same height. Individuals in Site B imitated the same pattern of horizontal arboreal plane display among the group. Discussion The marking technique we employed proved successful. The use of bands to mark lizards permitted identification of individuals from several meters away and throughout the entire study period because the bands remained in place the entire time. The bands did not reflect sunlight and did not dislodge with shedding of the skin. Further- more, the presence of the bands did not appear to in- crease predation vulnerability since the bands were thin and somewhat covered by the hind limbs. This method amphibian-reptile-conservation.org Figure 14. Ceratophora tennentii in Tangappuwa, Dumbara (Knuckles World heritage), Sri Lanka. can be used as a temporary, noninvasive marking tech- nique for other behavioral studies of lizards, instead of the traditionally-used toe clipping, which injures lizards and can alter their behavior. Lyriocephalus scutatus showed clear territorial main- tenance and site fidelity behaviors at the two study sites at Gannoruwa Forest Reserve. The territorial behavior of L. scutatus is a daily- synchronized behavior, initiating with a morning display session followed by ground Walking, and in the evening another display session. Behaviors in- cluded in territorial maintenance and site fidelity include Body -lift, Gular Sac Display, Head-bob, and Taii-wag. Observations and time budget analysis of the behav- iors of the studied lizards show that Body -lift. Head-bob, Gular Sac Display (shown only by males), and Tail-wag are important for site fidelity behavior. When lizards dis- play there is a regular order of behaviors (Jennings and Thompson 1999) that begin with Body-lift followed by Head-bob. While doing Head-bob the Gular Sac Display is also performed. Tail-wag is rare, but when performed it is normally after these previously mentioned behav- iors. While lizards are displaying they always hold their Body-lift for a long time and while performing other be- haviors simultaneously. Observed male lizards held their Body-lift from five to 30 minutes, and it is suspected that this might help them appear larger and help in mate at- traction. Gular Sac Display is only exhibited by males and may be important in sexual selection (Stuart-Fox and 110 October 2012 I Volume 5 I Number 2 I e56 Territorial behavior of Lyriocephalus scutatus Ord 2004). Body adjustments help lizards to locate one another. The upright position of display in L. scutatus, combined with their laterally placed movable eyes on the top of their head, enables them to see others in the group in such a way that lizards are able to distinguish other individuals by their side view. Many anurans exhibit synchronized calls known as “chorus” behavior (Narins 1992). Likewise, Lyriocepha- lus scutatus shows synchronized territorial maintenance behaviors within a particular group (i.e., individuals within the group display their territorial behaviors si- multaneously). When one particular individual starts to display, the other individuals in the same group eventu- ally start their display as well. Synchronized territorial maintenance behavior is important for the recognition of the territory of a particular individual relative to all other individuals in the group from one point of view. In general, among agamid lizards of Sri Lanka males are known to be territorial (Deraniyagala 1931, 1953; Manamendra-Arachchi 1998; de Silva et al. 2005) and they show territorial behaviors more than females and ju- veniles. Therefore, it is not entirely surprising that males of L. scutatus show Body-lift, Gular Sac Display, Head- bob, and Tail-wag whereas females do not. Adjustment and Still are not territorial maintenance behaviors be- cause all three reproductive groups show them in nearly equal frequencies, with males showing a slightly lower frequency than the others. Subadults showed the highest frequency of Walking among the observed behaviors. This may be due to the process of acquiring a permanent territory. Males were generally more active than females. This disparity be- tween the sexes suggests that Body-lift, Gular Sac Dis- play, Head-bob, and Tail-wag are vital territorial main- tenance behaviors since they occur most frequently in males. The three genera Lyriocephalus, Ceratophora, and Gonocephalus are consistently placed within the same clade of the acrodont lizard phylogeny (Macey et al. 2000). Ceratophora (Sri Lankan horned lizards) and Lyriocephalus are sister taxa (Schulte et al. 2002), while Gonocephalus, is the closest Southeast Asian relative of Lyriocephalus (Macey et al. 2000). The territorial be- havior of the endemic Leaf-nosed horned lizard {Cera- tophora tennentii) is somewhat similar to L. scutatus as observed in previous fieldwork (Fig. 14). They perform Body-lift and Head-bob but there is a clear difference in the way they hold the body in Body-lift, Ceratophora tennentii holds its body with a curvature of the spinal col- umn while positioning the legs in similar manner to that of L. scutatus. Observations on Gonocephalus sp. (Fig. 15) in Lambir Hills National Park, Sarawak, Malaysia show a similar territorial behavior to that of L. scutatus, with Body-lift and Gular Sac Display being performed in a similar manner. The results presented here show a large difference in the size of male and female territories. Females have Figure 15. Gonocephalus sp. in Lambir Hills National Park, Sarawak, Malaysia. larger home ranges compared to that of males, which may be due to highly territorial nature of males, and fe- males mainly moving about for feeding and mating. The female territories always overlapped with that of males, which suggests that a single male has access to one or two females. Subadults, on the other hand, have territo- ries that overlap with females and adult males. This may be due to them not being of breeding size and thus not a threat to the resident adult males. Territory size was not linked to the body size of the owner. The size of the territory might depend on the slope and other physical factors of the land, vegetation cover of the study area, structure of the forest, or human interfer- ence in the area. Males had their own defended tree and they do morning and evening displays while perched on that tree. On one occasion a female was found on one of the trees occupied by a male. This study shows that adult males of L. scutatus are highly territorial. Individual males maintain their ter- ritories, although their territories can overlap with fe- males and male subadults. Adults of arboreal Anolis spp. usually occupy vertical territories such as trees and walls. Since a lizard defends all of the area in which it is found, except perhaps resting and egg laying sites, terri- tory is almost equivalent with home range (Philibosian 1975; Jennings and Thompson 1999). Generally, a liz- ard spends the entire daylight period moving from one frequented perch site to another, often spending several minutes at a single site. A typical perch position is with the body vertical and head pointing toward the ground at October 2012 I Volume 5 I Number 2 I e56 amphibian-reptile-conservation.org 111 Bandara various angles. The primary activities within the territory include feeding, copulation, and defense, the latter usu- ally against members of the same species and sex, and of similar size. Adults tend to stay in one territory until death, while younger animals are more mobile. Juveniles are usually spatially separated from adults, perching on small rocks and low vegetation. Subadults are often tol- erated within adult territories and territories of males and females may overlap (Jennings and Thompson 1999). Conclusion The arboreal distribution of the individuals of L. scuta- tus in the same group is a significant behavior and may be novel. This behavior seems to permit the individuals within a group to spot all or most of the other individu- als at once, thus increasing the communication among individuals within the group. Further study should be performed to investigate this peculiar behavior of L. scu- tatus more thoroughly. Within the short period of time allowed for the present study, the arboreal distribution of individuals in same group is the foremost finding and it gives us evidence of the hidden eccentric behaviors that agamid lizards possess. Moreover, it may be that other territorial agamid lizards show a similar aerial horizontal distribution and synchronizing display as well. What is clear is that future studies on the behavior of agamid liz- ards of Sri Lanka are needed since much of their ecology remains unknown. Acknowledgments. — I thank Dr. (Mrs.) Suyama H. Meegaskumbura of Department of Zoology, Faculty of Science, University of Peradeniya for her kind guidance, advice and comments throughout the research. Moreover, I wish to thank Mr. Ruchira Somaweera, Mrs. Nilusha Somaweera, Mr. Nay ana wijetilake, and Mr. Kanishka Ukuwela for their constructive comments of selecting this research topic. I would also like to thank Chamara Jay aba Amarasinghe, Thilanka Ranathunge, Chathura Ekanayake, Kasun Munasinghe, Sandun Nalaka, Lahiru Malshan, and the members of Youth Exploration Society, Royal Botanic Gardens Peradeniya for their comments and field assistance. Special thanks go to Mr. Ruchira Somaweera Mr. Mendis Wickramasinghe, Mr. Kanishka Ukuwela and Mr. Sameera Karunarathna, Mr. Pradeep Samarawickrema, Mr. Samitha Harishchandra for pro- viding the necessary literature, for valuable critique and review of the manuscript, and for personal comments. I am also thankful to the villagers of Yatihalagala for their kind cooperation during the field studies and I gratefully acknowledge the comments by the anonymous review- ers. Literature cited Bahir MM, Surasinghe TD. 2005. A conservation assess- ment of the Sri Lankan Agamidae (Reptilia: Sauria). In: Contribution to Biodiversity Exploration and Re- search in Sri Lanka. Editors, Yeo DCJ, Ng PKL, Pe- hiyagoda R. The Raffles Bulletin of Zoology, Supple- ment 12:407-412. Bambaradeniya CNB, Samarawickrema PK, Ranawana KB. 1997. Some observations on the natural history of Lyriocephalus scutatus (Linnaeus, 1776) (Reptilia: Agamidae). Lyriocephalus 3(l):25-28. Deraniyagala PEP. 1931. Some Ceylon Lizards. Spoila Zelanica 16(2): 139-180. Deraniyagala PEP. 1953. A Colored Atlas of some Ver- tebrates from Ceylon. 2; Tetrapod Reptiles. Ceylon Government. Press. 35 plates -i- 101 p. De Silva A, Bauer A, Goonawardana S, Drake J, Na- thanael S, Chandrarathne R, Somathilaka S. 2005. Status of the agamid in the Knuckles Massif with special reference to Calotus liocephalus and Cophotis ceylanica. Lyriochephalus 06:43-52. Jennings WB, Thompson G. 1999. Territorial behavior in the Australian scincid lizard Ctenotus fallens. Herpe- tologica 55(3):352-361. Karunarathna DMSS, Amarasinghe AAT. 2008. Notes on the territorial behaviour of Otocryptis wiegmanni Wagler, 1830 (Reptilia: Agamidae: Draconinae). Her- petotropicos 4(2):79-83. Macey JR, Schulte II, Larson A, Ananjeva NB, Wang Y, Pethiyagoda R, Rastegar-Pouyani N, Papenfuss TJ. 2000. Evaluating trans-tethys migration: An example using acrodont lizard phylogenetics. Systematic Biol- ogy 49(2):233-256. Manamendra-Arachchi K. 1998. Gecko. Sri Lanka Na- ture l(l):45-55. Narins PM. 1992. Evolution of anuran chorus behavior: Neural and behavioral constraints. The American Nat- uralist 139:90-104. Philibosian R. 1975. Territorial behavior and population regulation in the lizards, Anolis acutus and A. cris- tatellus. Copeia 1975(3):428-444. Samarawickrama VAMPK, Ranawana KB, Rajapaksha DRNS, Ananjeva NB, Orlov LN, Ranasinghe JMAS, Samarawickrama VAR 2006. A new species of the ge- nus Cophotis (Squamata: Agamidae) from Sri Lanka. Russian Journal of Herpetology 13(3):207-214. Schulte JA II, Macey R, Pethiyagoda R, Larson A. 2002. Rostral horn evolution among agamid lizards of the genus Ceratophora endemic to Sri Lanka. Molecular Phylogenetics and Evolution 22(1): 11 1-1 17. Smith MA. 1935. The Launa of British India, including Ceylon and Burma. Reptilia and Amphibia. Volume II. - Sauria. Taylor and Francis, London, United Kin- dom. amphibian-reptile-conservation.org 112 October 2012 I Volume 5 I Number 2 I e56 Territorial behavior of Lyriocephalus scutatus Somaweera R, Somaweera N. 2009. Lizards of Sri Lan- ka: A Colour Guide with Field Keys. Chaimaira Pub- lications, Germany. 303 p. Stuart-Fox DM, Ord TJ. 2004. Sexual selection, natural selection and the evolution of dimorphic coloration and ornamentation in agamid lizards. Proceedings of the Royal Society B: Biological Sciences 271:2249- 2255. Wickramasinghe LJM. 2006. A new species of Cnemas- pis (Sauria: Gekkonidae) from Sri Lanka. Zootaxa 1369:19-33. Received: 29 May 2011 Accepted: 11 October 2012 Published: 27 October 2012 Imesh Nuwan Bandara obtained a Bachelor of Science (B.Sc.) degree specializing in zoology from the University of Peradeniya, Sri Lanka. Despite his love for nature and animals beginning in his early childhood, his “scientific” exploration of biodiversity began with him joining the Youth Exploration Society of Sri Lanka (Y.E.S.) in late 1990s. Since then he has been involved in multitude of nature-related activities, especially with regard to Sri Lankan unique fauna (his field experience as a freelance researcher/biologist primarily covers the fauna of Sri Lanka). Imesh is keen on studying much of the land vertebrates and invertebrates, their tax- onomy, life history, ecology, and conservation. He is also particularly interested in ethnobotany and cultural biodiversity of the island (Sri Lanka). Imesh has experience working in most of the Sri Lankan National Parks, Strict Nature Reserves, Protected Areas, other Eorest Reserves and rural village areas across the country be- ginning in 1998 through conducting, organizing, and consulting with biodiversity awareness programs in the conservation community. Imesh applies his knowledge of Sri Lankan herpetology to conserve some of the most threatened amphibian and reptile species of the island through various research and awareness programs. Imesh is a member and a former president of Y.E.S. His specific fields of research interest include: Ecosystem services, community based conservation, traditional agricultural practices, ethnobotany, local biodiversity, and behavioral ecology of herpetofauna and other wild fauna. amphibian-reptile-conservation.org 113 October 2012 I Volume 5 I Number 2 I e56 Copyright: © 2012 Dissanayake and Wellapuli-Arachchi.This is an open-access article distributed under the terms of the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License, which permits unre- stricted use for non-commercial and education purposes only provided the original author and source are credited. Amphibian & Reptiie Conservation 5(2) :1 14-1 24. Habitat preferences of the endemic shrub frog Pseudophilau- tus regius (Manamendra-Arachchi and Pethiyagoda 2005) at Mihintale Sanctuary, Sri Lanka ^Duminda S. B. Dissanayake and ^S. M. Wellapuli-Arachchi ^•^Department of Biological Sciences, Faculty of Applied Sciences, Rajarata University of Sri Lanka, Mihintale, SRI LANKA Abstract. — Mihintalae is situated in the dry zone of the North Central Province of Sri Lanka, at an elevation of 108 m, and is an under studied site of the habitat of the endemic shrub frog Pseu- dophiiautus regius. Six different habitat types which included forest edge, seasonal pond, rock, shrub, grassland, and home garden habitats were selected and systematically sampled to identify the habitat preference of P. regius. During the survey, a total of 143 P. regius individuals were count- ed. The highest percentage (53%) of individuals were recorded from the forest edge habitats, 23% from shrub land habitats, 20% from home gardens, and 2% from grassland and seasonal ponds. No individuals were found in the rocky areas. The number of observed individuals of Pseudophiiau- tus regius increased with the rainfall in forest habitats and simultaneously decreased in the home gardens. During the dry season the overall turnout of the number of individuals increased in home gardens. However, more extensive and systematic studies, over a longer period of time, are required to estimate the population size and document the fluctuation of P. regius and implement suitable conservation measures, if necessary. Key words. Pseudophilautus regius, habitat preference, Sri Lanka, Mihintale Sanctuary Citation: Dissanayake DSB, Wellapuli-Arachchi SM. 2012. Habitat preferences of the endemic shrub frog Pseudophilautus regius (Manamendra-Arach- chi and Pethiyagoda 2005) at Mihintale Sanctuary, Sri Lanka. Amphibian & Reptile Conservation 5(2):114-124 (e57). Introduction Sri Lanka is part of the Sri Lanka- Western Ghats bio- diversity hotspot with a rich herpetofaunal assemblage (Meegaskumbura et al. 2002; Bossuyt et al. 2004; Mee- gaskumbura et al. 2009; De Silva 2009; Meegaskumbura and Manamendra-Arachchi 2011). A total of 112 am- phibian species are known from Sri Lanka (De Silva et al 2005; Manamendra-Arachchi and Pethiyagoda 2005 and 2006; Meegaskumbura and Manamendra-Arachchi 2005; Meegaskumbura et al. 2010; Meegaskumbura and Manamendra-Arachchi 2011). Among the Sri Lankan amphibians, the most speciose family is the frog family Rhacophoridae. The Rhacophoridae consists of approxi- mately 321 species within two subfamilies and distrib- uted across a wide range of habitats in tropical Africa and south Asia, including India and Sri Lanka (Frost 2008; Li et al. 2008; Yu et al. 2008; Frost 2011). All the Sri Lank- an rhacophorids belong to the subfamily Rhacophorinae that contains three genera Pseudophilautus, Polypedates, and Taruga (Manamendra-Arachchi and Pethiyagoda 2005; Meegaskumburaet al. 2010; AmphibiaWeb 2011; Meegaskumbura and Manamendra-Arachchi 2011), of which Pseudophilautus is the most diverse with 68 spe- Correspondence. Email: ^duminda.rusl® gmail.com cies (Manamendra-Arachchi and Pethiyagoda 2005; Meegaskumbura and Manamendra-Arachchi 2005; Mee- gaskumbura et al. 2009; Meegaskumbura and Manamen- dra-Arachchi 2011). Amphibian diversity of Sri Lanka is directly influ- enced by climate, vegetation, topography, and geology, and its high rainfall and humidity provide ideal condi- tions for amphibians. The species richness of Pseu- dophilautus is greatest in the wet zone of Sri Lanka (Ma- namendra-Arachchi and Pethiyagoda 2005). The only two species of Pseudophilautus that have been reported hitherto from the dry zone of Sri Lanka are P. ferguso- nianus (Ahl 1927) and P. regius (Manamendra-Arachchi and Pethiyagoda 2005). Pseudophilautus regius is an en- demic species listed as Data Deflcient in the 2007 Red List of Threatened Fauna and Flora of Sri Lanka. This species is distributed in localized patches of the dry zone (De Silva et al. 2004; Manamendra-Arachchi and Pethi- yagoda 2005; Karunarathna and Amarasinghe 2007; Karunarathna et al. 2008; De Silva 2009) including the Mihinthale Sanctuary in the Anuradhapura District (Dis- sanayake et al. 2011). Pseudophilautus regius becomes active during the northeast monsoon and inter-monsoonal period (Bahir et amphibian-reptile-conservation.org 114 November 2012 I Volume 5 I Number 2 I e56 Dissanayake and Wellapuli-Arachchi al. 2005). However, very little is known about its breed- ing biology (Dubois 2004; Bahiret al. 2005), with the only report being that after amplexus, the female digs a small hole where she lays her eggs and then covers them with soil (Karunarathne and Amarasinghe 2007). Virtu- ally nothing is known about the population size, behav- ior, dispersal of non-breeding individuals, and habitat preferences of P. regius. This study was carried out to unravel the habitat preference of P. regius in the Mihin- tale Sanctuary. Methods and materials Study area Mihintale Sanctuary is located near the town of Mihin- tale (Anuradhapura District, North Central Province) in the dry zone of Sri Lanka. Annual rainfall in the area of Mihinthale is approximately 1,000-1,500 mm, with most of it occurring during the inter-monsoonal (October and November) and the north-east monsoonal (December un- til February) periods. The mean annual air temperature is 26 °C with a minimum of 19.5 °C and a maximum of 35 °C. The Mihintale Sanctuary is approximately 2,470 acres (999.6 ha) in extent with no proper demarcated boundaries (Fig. 1). Methods The study was carried out from October 2010 to March 2011, with the exception of February 2011. Quadrat sam- pling (Heinen 1992) in randomly selected points was per- formed within the Mihintale Sanctuary. A total of twenty- four 10 X 10 m quadrats were sampled at selected points in each habitat type. The habitat types sampled were: Forest Edge (FEH; Fig. 3), Seasonal Pond (SPH; Fig. 4), Rocky Area (RAH; Fig. 9), Shrub Area (SAH), Grass- land (GLH; Fig. 5), and Home Garden (HGH). Each habitat consisted of four fixed-quadrat sampling points. Field surveys were conducted from 1800 to 2200 hrs and each sampling site was visited twice a week. A minimum of four people were engaged in the sampling which in- volved sorting through all leaf litter and searching the branches, tree trunks, and logs within plots. Specimens were identified, photographed, and released at the site of capture. A structured data sheet was used to record data, including environment parameters such as air tempera- ture and relative humidity (RH), which were recorded using a thermometer (-20-100 °C, + 0.5 °C) and hygrom- eter (+ 4% RH at -r 77 °F within 10 to 90% RH + 5% RH at all other range) respectively. Results and discussion A total of 143 individuals of P regius (Fig. 2) were ob- served from six habitat types during the survey. The amphibian-reptile-conservation.org highest number was recorded from dry FEH (53%) (Fig. 3), followed by SAH (23%), HGH (20%), GLH (Fig. 5), and SPH (2%) (Fig. 4). No individuals were recorded from RAH during the survey period. These results suggest that the most preferred habitats of P regius are FEH, SAH, and HGH. Seasonal ponds provide good breeding sites for anurans (Conant and Col- lins 1991; Gibbs 2000), and according to Dissanayake et al. (2011) SPH had the highest percentage of amphib- ians recorded in the Mihinthale Sanctuary. However, we recorded few individuals in SPH. This could be because the habitat was surrounded by rocks with no moisture, no thick leaf litter layer (20 mm), or any significant canopy layer (over 70%). GLH was not covered with leaf litter and the area had a higher percentage of Imperata cylin- drica and Panicum maximum grasses, which might be a reason for the low number of individuals recorded in this habitat type, yet more than SPH. Most anurans are active during a confined period of time in the day or season (Peterson and Dorcas 1992). In many species, vocal advertisement represents the most energetically demanding behavior of males during the adult phase of the life cycle (Ryan 1983; Pough et al. 1992). Furthermore, the calls increase the probability of being exposed to predators. During the survey, most re- cordings of P. regius calling came from FEH and SAH. Stachytarpheta indica, Ageratum conyzoide, Clidemia hirta, Pterospermum suberifolium, Lantana camara, Zizyphus oenopila, Leucaena leucocephala. Acacia leu- cophloea, Drypetes sepiaria, Bauhinia racemosa, and Bridelia retusa were the abundant plant species in these two habitats. Average DBH in FEH was 16.26 cm, in- cluding trees with a DBH > 120 cm like Diospyrose eb- enum that, with small trees, provide a significant canopy layer (over 70%) and a thick leaf litter layer (20 mm). Therefore, FEH and SAH may provide the most pre- ferred habitats for P regius. The canopy cover (>70%) and a moist thick leaf litter layer (20 mm) are important to avoid desiccation and also to lay their direct develop- ing eggs (Bahir et al. 2005; Karunarathne and Amaras- inghe 2007). According to Menin et al. (2007) the contra- dictory relationship of anuran communities and the leaf litter layer can be related to different methods of quanti- fying litter characteristics such as volume, depth, and dry mass. On the other hand, relationships were found be- tween the depth of leaf litter in many studies on anurans in forests of Costa Rica (Lieberman 1986), Central Ama- zonia (Tocher et al. 1997), Uganda (Vonesh 2001), and the Southeast region of Brazil (Van Sluys et al. 2007). In the present study, analysis of rainfall patterns of the sampling locations revealed an increase in the number of observed individuals of P regius immediately after rain in FEH and SAH. This study is in agreement with previ- ous studies that seasonal variation of anuran populations is infiuenced by rainfall pattern (Das 1996; Weeraward- hena et al. 2004). Our data indicates that during the rainy period (monsoon and inter-monsoonal), the number of 115 November 2012 I Volume 5 I Number 2 I e56 Habitat preferences of the endemic shrub frog Pseudophilautus regius 1 200 m 1 1 500 ft 1 Figure 1. Map of study area. Human settlment and less forest area Forest area Grasslands Figure 2. Pseudophilautus regius (mature male). amphibian-reptile-conservation.org 116 November 2012 I Volume 5 I Number 2 I e56 Dissanayake and Wellapuli-Arachchi Figure 3. View of Forest Edge Habitat (FEH). Figure 4. View of Seasonal Pond Habitat (SPH). amphibian-reptile-conservation.org 117 November 2012 I Volume 5 I Number 2 I e56 Habitat preferences of the endemic shrub frog Pseudophilautus regius Figure 5. View of Grassland Habitat (GLH). Home gardens Grasslands Rocky areas Seasonal Forest edges Shrub lands Habitats Figure 6. Comparison of the percentage of Pseudophilautus regius found in each habitat type. amphibian-reptile-conservation.org 118 November 2012 I Volume 5 I Number 2 I e56 Dissanayake and Wellapuli-Arachchi Figure 7. Average rainfall (mm) from October 2010 to March 2011 at the Mihintale Sanctuary, indicating Forest Edge Habitat (FEH). Figure 8. Average rainfall (mm) from October 2010 to March 2011 at the Mihintale Sanctuary, indicating Home Garden Habitat (HGH). amphibian-reptile-conservation.org 119 November 2012 I Volume 5 I Number 2 I e56 Habitat preferences of the endemic shrub frog Pseudophilautus regius Figure 9. View of Rocky Area Habitat (RAH). Figure 10. Inside forest: Dry mixed evergreen vegetation with good leaf litter. amphibian-reptile-conservation.org 120 November 2012 I Volume 5 I Number 2 I e56 Dissanayake and Wellapuli-Arachchi individuals of P. regius increase in FEH (Fig. 7). Howev- er, our study was not conducted in February, although it rained in that month. This study is also in agreement with a study conducted in Madagascar where all amphibian species were edge-avoiders in the dry season but showed different patterns during the wet season (Lehtinen et al. 2003). In the dry months (October and March) however, the percentage of the number of individuals of P. regius were higher in HGH than in the rainy season (Novem- ber-January) (Fig. 8). This could be because HGH pro- vide various human modified microhabitats that attract frog species like P regius. A high number of individuals were observed near garden water taps and also near bath- rooms. This may be because during the dry season forest litter and soil dry-up, although some moisture remains around water taps due to dispersal of water during usage or due to leakages. However, this observation does not indicate that P. regius is solely found in disturbed habi- tats, and could be because this study was conducted for a short time period. Further research conducted at least for a year could reveal possible relationships with relative humidity Conclusions and recommendations The habitat type most preferred by P regius is Forest Edge Habitat (53%), whereas Rocky Area Habitat was not. The present study also demonstrates that Home Gar- den Habitat might provide suitable habitats during the dry season. Additional studies are needed using differ- ent sampling methods coupled with behavioral studies to determine the distribution of P. regius across the forest habitat and through home garden during the dry season. It was observed that villagers used Mihintale Sanctuary for daily activities including the forest edge for collecting firewood. Furthermore, some residents on the sanctuary boundary disturb the shrubs. These activities can have an adverse effect on the population of P regius. We also saw garbage accumulation in the sanctuary (Fig. 11), which may affect the breeding grounds as it pollutes the for- est fioor. We strongly suggest that management authori- ties take necessary steps to minimize and mitigate these adverse impacts in order to conserve the habitat of this endemic shrub frog. Long-term monitoring programs should be conducted to estimate the population fiuctua- tion and implement suitable conservation measures if necessary. Acknowledgments. — We are particularly grateful to Dr. Shirani Nathanael (Faculty of Applied Sciences, Rajarata University of Sri Lanka) and Mr. L. J. Mendis Wickramasingha (Herpetological Foundation, Sri Lanka) for supervision, unfailing encouragement, guidance, con- structive, but calm criticism and moral support to carry out our research. We wish to thank Mr. Niwanthaka San- Figure 11. Garbage accumulation in Mihintale Sanctuary. amphibian-reptile-conservation.org 121 November 2012 I Volume 5 I Number 2 I e56 Habitat preferences of the endemic shrub frog Pseudophilautus regius jeewa Thennakoon, Mr. Shiran Fernando, Mr. Jeevan Priyankara Karunarathna, Mr. Chathura Sandamal, Mr. Asela Dinushan, and Mr. Dushan Dharshanfor their sup- port with field work. Dr. T. V. Sundarabarathy and Dr. S. Wickramasinghe (Department of Biological Sciences, Faculty of Applied Sciences, Rajarata University of Sri Lanka) for their support in making this study a success. Mr. Dinidu Hewage assisted in developing the maps. We would like to thank the lUCN Librarian and D. M. S. Suranjan Karunarathna for providing literature and the Director General of the Meteorological Department for providing the relevant meteorological data to achieve our target. We also appreciate the valuable support provid- ed by K. G. D. de Aabeysinghe. Finally, we also like to thank Craig Hassapakis (ARC) who helped us in diverse ways to enrich this work. Literature cited Ahl E. 1927. Zur Systematik der asiatischen Arten der Froschgattung Rhacophorus. Sitzungsberichte der Gesellschaft Naturforschender Freunde zu (Berlin) 15:35-47. Bahir MM, Meegaskumbura M, Manamendra-Arachchi K, Schneider CJ, Pethiyagoda R. 2005. Reproduction and terrestrial direct development in Sri Lankan shrub frogs (Ranidae: Rhacophorinae: PhUatus). The Raffles Bulletin of Zoology (Supplement) 12:339-350. Bossuyt, F, Meegaskumbura M, Beenaerts N, Gower DJ, Pethiyagoda R, Roelants K, Mannaert A, Wilkinson M, Bahir MM, Manamendra-Arachchi K, Ng PKL, Schneider CJ, Oommen OV, Milinkovitch MC. 2004. Local endemism within the Western Ghats - Sri Lan- ka biodiversity hotspot. Science 306(5695):479-481. Conant R, Collins JT. 1991. Reptiles and Amphibians of Eastern and Central North America. Houghton Mif- flin, Boston, Massachusetts, USA. 608 p. Das I. 1996. Resource use and foraging tactics in a south Indian amphibian community. Journal of South Asian Natural History 2(1): 1-30. De Silva A. 2009. Amphibians of Sri Lanka: A Photo- graphic Guide to Common Frogs, Toads and Caeci- lians. Published by author. Creative Printers and De- signers, Kandy, Sri Lanka. 250 p. 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Why are there so many frog species in Sri Lanka? Alytes 22(l&2):19-37. Frost DR. 2008. Amphibian Species of the World: An Online Reference. Version 5.2 (15 July 2008). Ameri- can Museum of Natural History, New York, New York, USA. [Online]. Available: research.amnh.org/ herpetology/amphibia/php [Accessed: 26 September 2012 ]. Frost DR. 2011. Amphibian Species of the World: An Online Reference. Version 5.5. American Museum of Natural History, New York, New York, USA. [On- line]. Available: research.amnh.org/herpetology/am- phibia/php [Accessed:26 September 2012]. Gibbs JR 2000. Wetland loss and biodiversity conserva- tion. Conservation Biology 14(1):314-317. Heinen JH. 1992. Comparisons of the leaf litter herpe- tofauna in abandoned cacao plantations and primary rain forest in Costa Rica: Some implications for fau- nal restoration. Biotropica 24(3):43 1-439. Karunarathna DMSS, Amarasinghe AAT. 2007. 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Ecology of the leaf litter herpeto- fauna of an ecotropical rain forest: La Selva, Costa Rica. Acta Zoologica Mexicana (Nueva Serie) 15:1- 71. Manamendra-Arachchi K, Pethiyagoda R. 2005. The Sri Lankan shrub frog of the genus Philautus Gistel, 1848 (Ranidae: Rhacophorinae), with description of 27 new species. The Raffles Bulletin of Zoology (Supplement) 12:163-303. Meegaskumbura M, Bossuyt F, Pethiyagoda R, Mana- mendra-Arachchi K, Bahir MM, Milinkovitch MC, Schneider CJ. 2002. Sri Lanka: An amphibian hotspot. Science 298(5592):379. Meegaskumbura M, Manamnra-Arachchi K. 2005. De- scription of eight new species of shrub frogs (Rani- dae: Rhacophorinae: Philautus) from Sri Lanka. The Raffles Bulletin of Zoology (Supplement) 12:305-338. Meegaskumbura M, Manamendra-Arachchi K, Pethiya- goda R. 2009. Two new species of shrub frogs (Rha- cophoridae: Philautus) from the lowlands of Sri Lan- ka. Zootaxa 2122:51-68. Meegaskumbura M, Manamendra-Arachchi K. 2011 Two new species of shrub frogs (Rhacophoridae: Philautus) from Sri Lanka. Zootaxa 2747:1-18. Meegaskumbura M, Meegaskumbura S, Bowatte G, Manamendra-Arachchi K, Pethiyagoda R, Hanken J, Schneider CJ. 2010. Taruga (Anura: Rhacophoridae), A new genus of foam-nesting tree frogs endemic to Sri Lanka. Ceylon Journal of Science (Biological Sci- ences) 39(2):75-94. Menin M, Lima AR, Magnusson WE, Waldez F. 2007. Topographic and edaphic effects on the distribution of terrestrially reproducing anurans in Central Ama- zonia: Mesoscale spatial patterns. Journal of Tropical Ecology 17(2): 86-91. Peterson CR, Dorcas ME. 1992. The use of automated data acquisition techniques in monitoring amphibian and reptile populations. In: Wildlife 2001: Popula- tions. Editors, McCullough DR, Barrett RH. Elsevier Scientific Publishers LTD, Barking, Essex, England. 369-378. Rough EH, Magnusson WE, Ryan MJ, Wells KD, Taigen TL.1992. Behavioral energetics. In: Environmental Physiology of the Amphibians. Editors, Feder ME, Burggren WW. University of Chicago Press, Chicago, Illinois, USA. 395-436. Ryan MJ.1983. Sexual selection and communication in a Neotropical frog, Physalaemuspustulosus. Evolution 37(2):261-272. Tocher MD, Gascon C, Zimmerman BL. 1997. Fragmen- tation effects on a Central Amazonian frog commu- nity: A ten-year study. In: Tropical Eorest Remnants: Ecology, Management and Conservation of Erag- mented Communities. Editors, Laurance WE, Bier- regaard RO. University of Chicago Press, Chicago, Illinois, USA. 124-137. VanSluys M, Vrcibradic D, Alves MAS, Bergallo HG, Rocha CFD. 2007. Ecological parameters of the leaf- litter frog community of an Atlantic Rainforest area at Ilha Grande, Rio de Janeiro state, Brazil. Austral Ecology 32(3):254-260. Vonesh JR. 2001. Patterns of richness and abundance in a tropical African leaf-litter herpetofauna. Biotropica 33(3):502-510. Weerawardhena S, Amarasinghae US, Kotagama SW. 2004. Activity pattern and environmental variation of microhabitats of the six-toed green frog Euphlyc- tis hexadactylus Lesson 1834 (Anura-Ranidae) in Sri Lanka. Lyriocephalus 5(1&2):111-129. amphibian-reptile-conservation.org 123 November 2012 | Volume 5 | Number 2 | e56 Habitat preferences of the endemic shrub frog Pseudophilautus regius Yu G, Rao D, Yang J, Zhang M. 2008. Phylogenetic re- lationships among Rhacophorinae (Rhacophoridae, Anura, Amphibia), with an emphasis on the Chinese species. Zoological Journal of the Linnean Society 153(4):733-749. Received: 26 January 2012 Accepted: 30 April 2012 Published: 2 November 2012 Duminda S. B Dissanayake is a Sri Lankan undergraduate student pursuing a B.Sc. (Special) degree in the Department of Biological Sciences, Faculty of Applied Sciences, Rajarata University of Sri Lanka, Mihintale. Duminda began his career in wildlife research in 2009 with a focus on ecology and behavior of amphibians, reptiles, and birds. He is keen on spending his leisure time photographing wildlife and lending his support to wildlife conservation. Supun Mindika Wellappuli-Arachchi is an undergraduate student pursuing a B.Sc. (Special) degree in Fish- eries and Aquaculture, Department of Biological Sciences, Faculty of Applied Sciences, Rajarata University of Sri Lanka, Mihintale. His passion focused on nature from a very young age which subsequently led him in 2007 to a career in wildlife research (fish biology). His area of interest has now widened to include ecology and behavior of amphibians, reptiles, and birds as well and wildlife photography. He is dedicated to conserva- tion of biodiversity of Sri Lanka. amphibian-reptile-conservation.org 124 November 2012 I Volume 5 I Number 2 I e56 CONTENTS Administration, journal information (Instructions to Authors), and copyright notice Inside front cover Peter Janzen and Malaka Bopage — ^The herpetofauna of a small and unprotected patch of tropical rainforest in Morningside, Sri Lanka I Krishan Ariyasiri, Gayan Bowatte, Udeni Menike, Suyama Meegaskumbura, and Madhava Meegaskum- BURA — Predator-induced plasticity in tadpoles of Polypedates cruciger (Anura: Rhacophoridae) 14 Gayan Bowatte and Madhava Meegaskumbura — Morphology and ecology of Microhyla rubra (Anura: Mi- crohylidae) tadpoles from Sri Lanka 22 Walter R. Erdelen — Conservation of biodiversity in a hotspot: Sri Lanka’s amphibians and reptiles 33 Indika Peabotuwage, I. Nuwan Bandara, Dinal Samarasinghe, Nirmala Perera, Majintha Madawala, Chamara Amarasinghe, H. K. Dushantha Kandambi, and D. M. S. Suranjan Karunarathna — Range extension for Duttaphrynus kotagamai (Amphibia: Bufonidae) and a preliminary checklist of herpe- tofauna from the Uda Maliboda Trail in Samanala Nature Reserve, Sri Lanka 52 W. Madhava S. Botejue and Jayantha Wattavidanage — Herpetofaunal diversity and distribution in Kalugala proposed forest reserve, Western province of Sri Lanka 65 V. A. M. P K. Samarawickrama, D. R. N. S. Samarawickrama, and Shalika Kumburegama — Herpetofauna in the Kaluganga upper catchment of the Knuckles Forest Reserve, Sri Lanka 81 A. A. Thasun Amarasinghe, Franz Tiedemann, and D. M. S. Suranjan Karunarathna — Calotes nigrilabris Peters, 1860 (Reptilia: Agamidae: Draconinae): a threatened highland agamid lizard in Sri Fanka 90 Imesh Nuwan Bandara — ^Territorial and site fidelity behavior of Lyriocephalus scutatus (Agamidae: Draconinae) in Sri Lanka lOI Duminda S. B. Dissanayake and S. M. Wellapuli-Arachchi — Habitat preferences of the endemic shrub frog Pseudophilautus re gins (Manamendra-Arachchi and Pethiyagoda 2005) at Mihintale Sanctuary, Sri Lanka. . 114 Table of Contents Back cover volume 5 2012 NUMBER 2