Taprobanica (2009) Vol. 1. No. 1. Pages 01-77.

EDITORIAL

TAPROBANICA VOL. 01 No. 01

AN INTRODUCTION TO TAPROBANICA

Greek astronomer Ptolemy compiled a map in the 2nd century labeled the Island “Taprobane”. Arab

seafarers called it “Serendib”. From the 1400s, Europeans identified that island as “Seylan”, which was

later anglicized to “Ceylon”. In the 1500s Portugal and Spain established their dominance in the maritime

trade of South and Southeast Asia. In the 1600s the Dutch emerged as the dominant colonial power in the

region, followed in the 1700s by the British, then Independence Island after 1948 “Sri Lanka”. The island

is situated in the Indian Ocean, south west of the Indian peninsula, between 79o 39’ and 81

o 53’ Eastern

longitudes and 5o 54’ and 9

o 52’ Northern latitudes, and has a total land area of 65,610 km

2. “Taprobane”

was a historical name for Sri Lanka and we use the name “Taprobanica” for our society as well as for our

journal. Taprobanica Nature Conservation Society (TNCS) is an independent, non-governmental, non-

profit oriented voluntary organization dedicated to promote research, public awareness and nature

conservation and it was established in 2007.

According to the present literature information on behaviors, feeding and breeding habits, population

dynamics, ecology and threats are not properly understood for most of the species in Sri Lanka. Still a

considerable number of species to be explored and the taxonomic status to be classified systematically.

Therefore researches on the above aspects of Sri Lankan fauna are essential for the planning,

implementation of proper “scientific conservation” and management programs which should be improved.

Following the decline in the wake of independence from British in 1948, the past decade has seen an

acceleration of the rate of the biodiversity exploration in Sri Lanka. While these works have been mostly

of high quality, several papers has been published by local scientists unfamiliar with the science of

taxonomy and the procedures of the International Code of Zoological Nomenclature (ICZN), resulting in

the establishment of several unnecessary new names and invalid and erroneous information on Sri Lankan

fauna. In such a situation, we have initiated publishing the Taprobanica, The Journal of South Asian

Biodiversity largely in order to provide the opportunity to scientists on the natural history of Sri Lanka and

Western Ghats to publish their results in a journal with reasonably high standards of production and

review.

Taprobanica publishes original research papers, taxonomic descriptions, notes, observations, essays,

opinions and short communications. The Taprobanica is published in one volume comprising two

fascicles each year, starting with the first issue; volume 01 and number 01 in April 2009. The number of

fascicles/supplements/special issues/occasional papers may be increased to six depending on the volume

of contributions. Each fascicle of the journal will comprise 75-125 pages in perfect-bound double column

format, printed on high quality mat paper. High quality colour and black & white illustrations are

encouraged, and will be accepted with no charge forpages and illustrations. The journal is expected to be

circulated among leading conservation institutions and individuals in the field of ecology, biology and

natural history.

The journal’s geographic range extends to South Asia with special attention to Sri Lanka and Western

Ghats biodiversity hotspot. We would like to provide space in equal parts to ecology, geology, biology,

ethology, conservation breeding, conservation, evolution, morphology, physiology and systematics, but

excludes techniques or sampling methods. This first issue deals mostly with Sri Lankan herpetofauna,

because there is an increasing interest on herpetofauna than any other field in the subcontinent, which is

one of the most populous biodiversity hotspots identified for conservation.

A. A. Thasun Amarasinghe

Managing Editor: Taprobanica, the journal of South Asian Biodiversity

April 06th, 2009

TAPROBANICA, ISSN 1800-427X. April, 2009. Vol. 01, No. 01: pp. 1.

© Taprobanica Nature Conservation Society, 146, Kendalanda, Homagama, Sri Lanka.

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AMARASINGHE ET AL., 2009

TAPROBANICA VOL. 01: NO. 01

THE ORIGINAL DESCRIPTIONS AND FIGURES OF SRI LANKAN AGAMID LIZARDS (SQUAMATA: AGAMIDAE) OF THE 18TH AND 19TH CENTURIES

Submitted: 20 January 2009, Accepted: 27 January 2009

A. A. Thasun Amarasinghe 1,2, Ulrich Manthey 3, Edi Stöckli 4, Ivan Ineich 5, Sven O Kullander 6, Franz Tiedemann 7, Colin McCarthy 8 and Dinesh E. Gabadage 1

1 Taprobanica Nature Conservation Society, 146, Kendalanda, Homagama, Sri Lanka. 3 Society for Southeast Asian Herpetology, Kindelbergweg 15, D-12 249 Berlin, Germany. 4 Naturhistorisches Museum Basel, Augustinergasse 2, CH-4001 Basel, Switzerland. 5 Muséum national d’Histoire naturelle, Département Systématique et Evolution, USM 5502 (Reptiles), CP 30, F-75005 Paris, France. 6 Swedish Museum of Natural History, Department of Vertebrate Zoology, SE-104 05, Stockholm, Sweden. 7 Naturhistorisches Museum Wien, Herpetologische Sammlung, Burgring 7, A-1010 Vienna, Austria. 8 The Natural History Museum, Cromwell Road, London SW7 5BD, United Kingdom.

Corresponding authors: 2 [email protected], 6 [email protected] Abstract Eighteen species of agamid lizards are recognized from Sri Lanka, representing one subfamily: Draconinae. Thirteen of these species were described in the period 1758-1887. The early descriptions of agamid lizards were very brief and often written in languages other than English. Original descriptions and images of type specimens are provided for Calotes calotes (Linnaeus, 1758), C. versicolor (Daudin, 1802), C. nigrilabris Peters, 1860, C. liocephalus Günther, 1872, C. liolepis Boulenger, 1885, C. ceylonensis Müller, 1887, Otocryptis wiegmanni Wagler, 1830, Ceratophora stoddartii Gray, 1835, C. tennentii Günther, 1861, C. aspera Günther, 1864, Lyriocephalus scutatus (Linnaeus, 1758), Cophotis ceylanica Peters, 1861, and Sitana ponticeriana Cuvier, 1829. Translations to English are provided of original descriptions that were published in Latin, French, and German. Key words: Original description, type specimen, Agamidae, Taxonomy, English translation, Sri Lanka

TAPROBANICA, ISSN 1800-427X. April, 2009. Vol. 01, No. 01: pp. 2-15, 4 pls. © Taprobanica Nature Conservation Society, 146, Kendalanda, Homagama, Sri Lanka.

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THE ORIGINAL DESCRIPTIONS AND FIGURES OF SRI LANKAN AGAMID LIZARDS OF THE 18TH AND 19TH CENTURIES


Introduction

Sri Lanka is a biodiversity hotspot together with the Western Ghats of Southern India and the area is rich in herpetofaunal assemblages (Bossuyt et al., 2004; Meegaskumbura et al., 2002; Myers et al., 2000). Eighteen species of agamid lizards, family Agamidae, have been recognized from Sri Lanka and fifteen (83%) of them are endemic to the island (Bahir & Surasinghe, 2005; de Silva, 2006; Manamendra-Arachchi et al., 2006), representing a single subfamily, the Draconinae (Macey et al., 2000). The genus Calotes is represented by seven species (Bahir & Maduwage, 2005). The genera Ceratophora, Lyriocephalus and Cophotis are endemic to Sri Lanka (Manamendra-Arachchi et al., 2006), with five species of Ceratophora (Pethiyagoda & Manamendra-Arachchi, 1998), two species of Cophotis (Manamendra-Arachchi et al., 2006), and one species of Lyriocephalus (de Silva, 2006). The other two genera are Sitana with one species (de Silva, 2006), and Otocryptis with two species (Bahir & Silva, 2005).

Thirteen of the Sri Lankan agamid species were formally described over a period of 130 years between 1758 and 1887, starting with Calotes calotes (Linnaeus, 1758) from Zeylona (present day Sri Lanka), and ending with the description of Calotes ceylonensis Müller, 1887. Only five of those original descriptions were written in English (Boulenger, 1885; Gray, 1833-35; Günther, 1861, 1864, 1872). The others appeared in French (Daudin, 1802; Cuvier, 1829) or German (Müller, 1887; Peters, 1860, 1861) or Latin (Linnaeus, 1758; Wagler, 1830). After more than a century of the description of Calotes ceylonensis, two new species, Ceratophora karu and Ceratophora erdeleni, were described (Pethiyagoda & Manamendra-Arachchi, 1998), followed by Calotes desilvai Bahir & Maduwage, 2005; Otocryptis nigristigma Bahir & Silva, 2005 and Cophotis dumbarae Manamendra-Arachchi et al., 2006. Most of the early descriptions were very brief compared to the recent descriptions. Several were also written in languages, familiar to scientists of those days but now much less familiar to the taxonomists. Original descriptions of these agamids were published in books and periodicals that today are difficult to obtain other than possibly from major libraries. In this paper we retyped those complete original descriptions with the English translations, in the hope that they will be useful for taxonomists interested in the Agamid lizards of Sri Lanka.

Materials and Methods This work is mainly based on original descriptions and type specimens. It contains copies of the original descriptions of all the agamid lizards of Sri Lanka published in the 18th and 19th centuries. We attempted to retain all pertinent text and figures, but format changes were necessary. Some illustrations have been reduced and pagination has changed; therefore, some text references will not necessarily refer to specific pages in this document. All the figures of the original descriptions and the photographs of each type specimen are shown in each plate accompanied with the referred species name (Plates 1-4). The English translation of each species description is presented below the original description when it written in another language. The material examined are from the BMNH, Natural History Museum, London, UK; MNHN, Muséum national d’Histoire naturelle, Paris, France; NMB, Naturhistorisches Museum, Basel, Switzerland; NMW, Naturhistorisches Museum Wien, Vienna, Austria; NRM, Swedish Museum of Natural History, Stockholm, Sweden; UUZM, Museum of Evolution, Uppsala, Sweden; and ZMB, Zoologisches Museum der Humboldt-Universität zu Berlin, Germany. All the photographs of type specimen are displayed with the photographers’ initials; Franz Tiedemann (FT), Colin McCarthy (CM), Ivan Ineich (II), Mark-Oliver Rödel (MO), Edi Stöckli (ES), Bodil Kajrup (BK) and Rainer Günther (RG).

Additional Information Lacerta calotes was described by Linnaeus (1758: 207) based on three literature references, viz. (1) the description of "Lacerta cauda longa, pedibus pentadactylis, dorso antice dentato, capite pone denticulato" in Linnaeus (1749: 289), which also includes references to “Lacerta ceilonica cærulea” in Seba (1734: 150, pl. 34, fig. 4), “Lacerta ceylonicus amphibius" in Seba (1734: 149, pl. 95, fig. 3), and “Lacerta ceilonica lemniscata …” in Seba (1734: 146, pl. 93, fig. 2); (2) the listing of Lacerta Calotes in Linnaeus (1754: 44), with references to “Lacerta cauda tereti longa ...” in Linnaeus (1749), "Systema Naturae 36 n. 13" (Linnaeus, 1748: 36), plus the Seba references in Linnaeus (1749), except that the figure given for “Lacerta ceilonica caerulea” is here pl. 34, fig. 4; and (3) Seba (1734: pl. 95, figs 3-4, pl. 93, fig. 2). Plate 93, fig. 2, and plate 95, figs. 3-4 in Seba (1734; 2006), indeed show an agamid lizard with

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bluish or blue-green vertical bars. There is no figure of Lacerta Calotes in Linnaeus (1754), in which work lizards are not illustrated. Lacerta Calotes is also one of several species of lizards in Linnaeus (1754) which is not described, but only listed with references. The specimen described in Linnaeus (1749) is currently catalogued as UUZM Reptilia 33 (Wallin, 2001), and that described by Linnaeus (1754) as NRM 106. Lönnberg (1896) listed the UUZM syntype and Andersson (1900) listed the NRM syntype as present in the respective collections. These two are thus the only surviving syntypes of Lacerta calotes Linnaeus, 1758, to our knowledge. The syntype series is made up of the two specimens in Sweden and the three illustrated by Seba. It is possible that NRM 106 and UUZM 33 represent specimens obtained from Seba's collection, but it is not possible to correlate the specimens using the data or figures in Seba (1734), as there are no obvious shared individual peculiarities available which would support specimen identification.Linnaeus (1749) description is based on a specimen contained in a donation from Prince Adolf Fredrik in 1745 to the Academy in Uppsala. The 1754 listing is based on a specimen in the collection of Adolf Fredrik, now King. There are no collection data preserved with the specimens, so all information regarding the origin of the specimens is that which can be deduced from Linnaeus's publications. This brings into question the type locality. Linnaeus (1749) did not provide locality data. Linnaeus (1754) gave the distribution as India. Linnaeus (1758) gave Zeylona [=present day Sri Lanka] for distribution. It may very well be that the distribution was derived from the information in Seba (1734), rather than associated with the specimen.

The types of Calotes versicolor (Daudin, 1802) have been considered to be present in the MNHN collections on several occasions (e.g., Kuhl, 1820; Zug et al., 2006) but several studies indicated that the types of Calotes versicolor were no more present in the MNHN collections (e.g., Brygoo, 1988; Wermuth, 1967). Duméril & Bibron (1837) stated that Daudin (1802) had only one juvenile specimen for his description. However Daudin clearly stated that he had two specimens. However Duméril & Bibron (1837) did not compare their specimens with the specimen of Daudin, thus probably the types were not in MNHN at that time. According to Duméril & Duméril (1851), there were two specimens of juveniles from East Indies

(Indes orientales) from the Seba’s collection, and they also mentioned that these could be the types of Daudin (but they are without collector’s name). According to the original description of Daudin (1802) both specimens examined might have been adults considering their body sizes, however Daudin only give the size for one specimen and did not mentioned the second. Thus the two specimens of Seba can not be the types of Daudin. In the original description the author stated that he recognized that species on Seba’s plate illustrating a lizard from Brazil and did not give their origin (Daudin, 1802). Pondicherry was sometimes considered as the type locality and Leschenault the type’s collector (e.g., Kuhl, 1820; Zug et al., 2006). However now the MNHN collection only includes one specimen (MNHN 2548) from Pondicherry given by Leschenault. This is a syntype of Agama tiedemanni Kuhl, 1820. Leschenault participated as botanist on the Terres Australes expedition of Captain Baudin which left from Le Havre (France) on 19 October 1800. He stopped at Timor when going back in 1803 due to illness. He explored Java and arrived back in France in 1807 only. He was not able to send material from Pondicherry to Daudin (description of Calotes versicolor in 1802) during this time. In May 1816 he embarked for India and only returned to France in 1822 (Bauchot et al., 1990; Lescure & Marty, 2000). However the type specimens of C. versicolor are no longer present in the MNHN collection, the illustration of Seba is not a type and it is a Brazilian animal. It could be the Amazonian iguanid Plica umbra. We consider that the only still available part of that type material is the specimen depicted on plate 44 of Daudin (1802) (see pl. 01, fig. 05) with the present evidence that the types have to be considered as missing from MNHN collections or lost shortly after their description; however the most realistic picture is that these types never were in MNHN collections.

One specimen of Calotes ceylonensis (NMB 3340), from “Kumbukan-aar (S.-O.-Ceylon)” is labelled as holotype in the NMB collections. The other specimen (NMB 3341), is from “N.-O. Provinz” and labeled as paratype. Kumbukan-aar probably refers to a river in the extreme southeast of Sri Lanka called Kumbukkan Oya on current maps (Karunarathna et al., 2009). The name Otocryptis becomes available from Wagler (1830), who indicated Wiegmann as the author (see Wagler, 1830). Wiegmann’s publication

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of the name however, although dated May 1830, was issued only in 1831 (Wiegmann, 1831). Wagler (1830) nowhere explicitly stated that his account was based on Wiegmann’s data, or on a communication by him, or on his then unpublished manuscript, and it is therefore Wagler who was responsible for the conditions making the name available, and who is thus author of the name (Bahir & Silva, 2005). The species name O. wiegmanni also becomes available from Wagler (1830), and has priority over O. bivittata Wiegmann, 1831. No type material was designated by Wagler for O.

wiegmanni, because Wagler (1830) made no reference to any material before him and attributed the genus name to Wiegmann (see Wagler, 1830). Although Wiegmann (1831) correctly noted the type locality of O. bivittata as Ceylon (= Sri Lanka), the type locality of O. wiegmanni specified by Wagler — America — is evidently a lapsus (Bahir & Silva, 2005). The label of the jar of the holotype of O. bivittata, ZMB 708, clearly states that the specimen is from Ceylon and mentions Bloch, presumably M. E. Bloch, for information on provenance see Bahir & Silva (2005).

Original descriptions of Sri Lankan Agamids, with English translations

Calotes calotes (Linnaeus, 1758) (Plate: 1) Linnaeus, C., Systema naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tomus I. Editio decima, reformata, 1758: 207. Material examined: One of two syntypes [note: the Seba specimens are also syntypes] Male (112.0 mm SVL); Cat. no. NRM 106; Loc. Zeylona: Asia (= Sri Lanka: Asia); Coll. Unknown; Date. Unknown (Fig. 1, 2).

Original description: Calotes. 29. L. [=Lacerta] cauda tereti longa, dorſo antice capiteque poſtice dentato. Amæn. Acad. I. p. 289. Muſ. Ad. Fr. I. p. 44. Seb. muſ. I. t. 95. f. 3, 4. (Fig. 3, 4)

t. 93. f. 2. Habitat in Aſia: Zeylona Corpus cæruleum ſquamis acutis, ſubtus ſtriatum. Spinæ dorſi lamellatæ.

English translation:

Calotes. 29. L. [=Lacerta] with long narrow tail, back anteriorly crested, posteriorly toothed. Amoenitates Academiae. I. p. 289. Museum Adolphi Friderici. I. p. 44. Seba‘s Museum. I. pl. 95. fig. 3, 4.

pl. 93. fig. 2. Habitat in Asia: Ceylon Body bluish, spiny scales, striated above. lamellate dorsal crest.

Calotes versicolor (Daudin, 1802) (Plate: 1) Daudin, F. M., Histoire Naturelle des Reptiles, III, 1802: 395-397, pl. XLIV. (Fig. 5) Material examined: Type lost. The specimen MNHN 2548, collected from Pondicherry by Leschenault, cannot be a syntype of C. versicolor Daudin 1802, because Leschenault visited Pondicherry only after 1816. According to Daudin’s original description (1802) one of his two syntypes measured 81 mm SVL, 202.5 mm tail length and 283.5 mm total length. Original description: L’AGAME ARLEQUINÉ, A DEUX RAIES (1). Voyez la planche XLIV de ce volume.

Ce reptile est très-semblable aux agames ondulé et hexagone, par la forme de ses diverses parties, et par celle de ses écailles; mais il est beaucoup plus remarquable que la plupart des animaux qui composent ce

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genre, soit à cause de sa taille svelte et de l’extrême longueur de sa queue, soit à cause des couleurs brillantes et rembrunies, dont le dessus de son corps est agréablement varié. Sa tête et son corps ont trois pouces de longueur, et sa queue a jusqu’à sept pouces et demi, ce qui fait une longueur totale de dix pouces six lignes. La tête est arrondie, presque quadrangulaire, un peu courte, munie derrière l’occiput d’écailles redressées en pointes, avec une très-petite crête épineuse, formée de petites écailles pointues, prenant sa naissance sur la nuque, et disparoissant ensuite peu à peu, à mesure qu’elle s’approche vers la base de la queue. La tête est blanchâtre uniforme en dessus, avec six traits bruns rayonnés autour des yeux, quatre traits bruns en travers sur le crâne, et un point brun sur l’occiput. Deux lignes longitudinales, blanches et droites, prennent leur origine derrière les yeux, et se prolongent ensuite sur chaque côté du dos jusques dessus la base de la queue. Dessus le corps on voit huit ou neuf bandes transversales, d’un brun plus foncé en devant, plus pâle en arrière, séparées par d’autres bandes d’un beau bleu clair et nacré. Ces bandes ne sont pas tranchées net sur les bords, mais elles paroissent au contraire s’y fondre entre elles et s’y méler: elles sont en outre séparées en trois portions, par les deux lignes longitudinales blanches qui sont sur les côtés du dos. La queue est deux fois et demie environ aussi longue que le reste de l’animal; et elle est très-amincie, sur-tout à son extrémité. J’ai trouvé deux individus de ce joli et brillant saurien dans la collection du muséum d’histoire naturelle de Paris: comme il paroît avoir parfaitement conservé ses couleurs dans l’esprit de vin où il est enfermé, je crois que la description que je viens d’en donner ne pourra manquer de satisfaire les naturalistes. Cet animal est assez bien gravé et fort mal enluminé dans l’ouvrage de Seba (tom. I, pl. XCII, fig. 1), sous le nom de lézard téjuguacu du Brésil: voyez ce que j’en ai dit précédemment à l’article de l’agame umbre. (1)

Agama versicolor; lineâ longitudinali albâ in utroque latere dorsi, tæniis transversis fuscis et lætè

cæruleis, caudâ duplò longiore.

Lacerta brasiliensis, tejuguacu. Seba, Thes. tom. I, pag. 144, pl. XCII, fig. I. (see pl. 1, Fig. 6).


Harlequin agama, with two bands (1). See the plate XLIV in this volume This reptile is very similar to the undulate [Agama undulata, Lacerta undulata Bosc – USA, Carolina] and hexagonal [Agama angulata, Lacerta angulata Rolander – Guyana] agamas by the shape of its different parts, and by that of its scales; but it is more remarkable than most of the animals making up this genus, for reason of its more slender size and the extreme length of its tail, or because of its bright and dull colours with which its back is nicely adorned. Its head and body are three [French] inches [=81 mm] in length, and its tail up to seven and a half inches [202.5 mm], making a total length of ten inches and six [French] lines [1 French line = 2.25 mm; 10 French inches + 6 French lines = 283.5 mm]. The head is rounded, nearly quadrangular, a little short, bearing scales modified into spines at the back of the occiput, with a very short spiny crest, formed of small spiny scales, beginning on the neck, and disappearing progressively when approaching the base of the tail. The head is uniform whitish above, with six brown stripes around the eyes, four brown stripes across the top of the head, and a brown spot on the occiput. Two longitudinal lines, white and straight, have their origin behind the eyes and continue extended on each side or the back to above the base of the tail. Above the body, one sees eight or nine transverse bands, of a darker brown in front, paler behind, separated by other bands made of a beautiful light pearly light blue. These bands are not clearly delimited at their edges, but they rather seem to diffuse between them and mix together: they are moreover separated into three divisions, by the two longitudinal white lines that run on the sides of the back. The tail is about two and a half times as long as the rest of the animal; and it is very narrowed, above all at its distal part. I found two specimens of this beautiful and brilliant saurian in the collection of the Paris natural history museum: as it seems to have conserved its colours perfectly in the spirits-of-wine in which it is preserved, I believe the description that I gave here will not fail to satisfy the naturalists. That animal is relatively well engraved and very badly illuminated in the work of Seba (tom. 1, pl. xcii, fig. 1), under the name of lizard téjuguacu of Brazil: see what I said previously about it in the part of the agame umbre. (1) Agama versicolor; longitudinal white lines on each side of the back, tawny brown and beautiful blue

transversal bands, tail two times as long [as body] Lacerta brasiliensis, tejuguacu. Seba, Thes. vol. I, pag. 144, pl. xcii, fig. 1.

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Calotes nigrilabris Peters, 1860 (Plate: 1) Peters, W. C. H., Monatsberichte der Königlichen Akademie der Wissenschaften zu Berlin, 1860: 183. Material examined: Holotype Male (99.8 mm SVL); Cat. no. NMW 23355 (see Tiedemann et al., 1994); Loc. Newera Ellia: Ceylon (= Nuwara Eliya: Sri Lanka); Coll. Unknown; Date. Unknown (Fig. 7, 8).

Original description:

Calotes (Bronchocele) nigrilabris n. sp.; oben olivenfarbig, unten hellgrün; die Schuppen des Kinnes und der Kehle an der Basis, die Lippenränder und die ganze Schläfengegend zwischen Augen und Trommelfell braunschwarz; die Gegend zwischen den Ohren und die Oberseite des Halses grünlichgelb. Ein Schuppenkamm von ähnlicher Gestalt wie bei Calotes ophiomachus erstreckt sich von dem Nacken bis zu der Basis des Schwanzes; ein Kamm von sieben verlängerten Schuppen jederseits über und hinter der Ohröffnung; eine sehr große mit kleinen Schuppen ausgekleidete Grube vor jeder Schulter. Sämmtliche Körper und Schwanzschuppen gekielt. Seitliche, mit ihren Kielen nach hinten und abwärts gerichtete Körperschuppen ungefähr halb so groß wie die mittleren Bauchschuppen. Schwanz- und Kehlschuppen etwa doppelt so groß wie die Bauchschuppen. Vierte Hinterzehe nur wenig länger als die dritte; der Nagel der dritten erreicht die Basis des Nagelgliedes der vierten Zehe. Schuppen in der Körpermitte in 47 Längsreihen. Schwanzschuppen in etwa 13 Längsreihen. Totallänge 0m, 40; Entfernung vom Schnauzenende bis zum After 0,096; Länge des Kopfes 0,035; der vordern Extremität 0,058; der hintern Extremität 0,085; der vierten Zehe 0,024; der dritten Zehe 0,020. Fundort: Newera Ellia; ein einziges Exemplar.


Calotes (Bronchocele) nigrilabris n. sp.; above olivaceous, below light green; the scales of the chin and the throat base, the labial margins and the whole temporal region between eyes and eardrum fuliginous; the region between the ears and the upper side of the neck greenish yellow. A scale ridge similar to the one of Calotes ophiomachus reaches from the nape to the tail base; a ridge composed of seven extended scales on each side above and behind the ear aperture; a very large pit lined with small scales in front of each shoulder. All the body and tail scales are carinate. Lateral scales with their ridges directed backwards and downwards about half the size of the middle abdominal scales. Tail and throat scales about twice the size of the abdominal scales. The fourth rear toe only a little longer compared to the third one; the nail of the third toe reaches up to the base of the nail bearing link of the fourth toe. Scales along the middle of the body arranged in 47 horizontal rows. Tail scales in about 13 horizontal rows. Total length 0,40m; distance from tail end to anus 0,096; head length 0,035; the front limb 0,058; the rear limb 0,085; the fourth toe 0,024; the third toe 0,020. Collecting locality: Newera Ellia; a single specimen.

Calotes liocephalus Günther, 1872 (Plate: 2) Günther, A., Annals and Magazine of Natural History, ser. 4, IX, 1872: 86. Material examined: Holotype Male (90.0 mm SVL); Cat. no. BMNH 1946.8.11.33; Loc. Peradeniya Distr., Ceylon (= Sri Lanka); Coll. G.H.K. Thwaites; Date. Unknown (Fig. 9, 10). Original description: No spines whatever on the side of the head. Dorsal crest composed of slender spines of moderate length on the neck, a low, merely serrated crest in the middle of the trunk, but reappearing in the sacral region as a short series of three or four spines. A very distinct fold in front of the shoulder, covered by granular scales. Gular sac very slightly developed. About forty-five series of scales round the middle of the trunk. Scales round the part of the tail in which the penis is hidden much the largest. Green, with irregular dark cross bands on the back. Upper side of the head marbled with dark green. A narrow green band from the eye to

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above the tympanum. Tail olive, with broad brown rings. Limbs with alternate lighter and darker green rings. One adult male is 15 inches long, the tail being 11 inches.

Calotes liolepis Boulenger, 1885 (Plate: 2) Boulenger, G. A., Catalogue of the Lizards in the British Museum, I, 1885: 326-327, pl. 25, fig. 2. (Fig. 11) Material examined: One of two syntypes Female (76.7 mm SVL); Cat. no. BMNH 69.7.24.2; Loc. Ceylon (= Sri Lanka); Coll. R.H. Barnes; Date. Unknown (Fig. 12, 13)


Upper head-scales smooth, imbricate, strongly enlarged on supra-orbital region; two distant spines on each side of the back of the head, between the ear and the nuchal crest; diameter of the tympanum nearly half that of the orbit. Gular sac not developed (♀); gular scales strongly keeled, as large as ventrals. A short oblique fold in front of the shoulder. Nuchal crest formed of narrow separated spines, the longest of which measure about the diameter of the tympanum; dorsal crest quite indistinct. Thirty-five or thirty-nine scales round the middle of the body; dorsal scales three times as large as ventrals, squarish, smooth, pointing backwards and downwards; ventral scales strongly keeled. The adpressed hind limb reaches hardly the tympanum; third and fourth fingers equal. Tail round. Pale olive, with indistinct brown transverse bands on the back; brown lines radiating from the eye, the lower reaching down to the lower lip.

Total length………………… 256 millim. Head…………………………. 21 ,, Width of head……………….. 13 ,, Body…………………………. 55 ,, Fore limb…………………….. 36 ,, Hind limb……………………. 51 ,, Tail…………………………. 180 ,,

Ceylon. a. ♀. Ceylon. B. H. Barnes, Esq. [P.]. b. Yg. Ceylon.

Calotes ceylonensis Müller, 1887 (Plate: 2) Müller, F., Verhandlungen der Naturforschenden Gesellschaft in Basel, 8 (2), 1887: 292-293, pl 3. (Fig. 14) Material examined: One of two syntypes Male (81.9 mm SVL); Cat. no. NMB 3340 (labelled as holotype); Loc. Kumbukan-aar: South East Ceylon (= Kumbukkan Ara ?: Sri Lanka); Coll. P. Sarasin & F. Sarasin; Date. 1886. (Fig. 15, 16)


Calotes mystaceus var. ceylonensis (an n. sp.?) (Taf. 3) Kumbukan-aar (S.-O.-Ceylon) und N.-O. Provinz Ceylon v. d. Hh. Sarasin. [2] (Hierzu Taf. III.) Die vorliegenden vorzüglich conservierten Stücke zeigen mehrere nicht unerhebliche Abweichungen von den bestehenden Beschreibungen. (DB., Gü. R. b. J., Boul. cat.). Schuppen der Kopfoberfläche glatt, ziegelig, jede am freien Rund mit einem Kranz von Körnchen besetzt, Schuppen der Supraorbitalregion beträchtlich grösser; über dem Trommelfell jederseits 2 Dornen distant. Trommelfell wenigstens ½ Durchmesser der orbita. – Kein Gularsack. – Kehlschuppen sehr stark gekielt, viel kleiner als Rückenschuppen (8:5). Vier bis fünf schwache Dornen auf dem Nacken; Rückenfirst ohne jede crista noch Denticulation. Rund um die Mitte des Körpers 60 Schuppen. Rückenschuppen kaum merklich gekielt, manche ganz glatt, 1 ½ mal so gross als Bauchschuppen. Bis zur Mitte des Körpers sind alle Schuppen aufwärts and rückwärts, auf der Hinterhälfte beinahe direct rückwärts gerichtet.

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Bauchschuppen mucronirt. Das angelegte hintere Glied reicht etwa bis zum hintern Augenrand. Vierter und fünfter Finger gleich lang. Vom Auge zum Trommelfell eine Reihe etwas grösserer Schuppen. Grünlich; über den Rücken 6 grosse sattelartige Querflecke, von denen die 3 vordern orangefarbenen unmittelbar aufeinander folgen und durch dunkle Linien abgegrenzt sind, die 3 hintern distanter, verschwommener und schwächer gefärbt sind. Auch quer über den Kopf ziehen mehrere helle Binden. Am Anfang des Nackenkammes ein schwarzer Doppelfleck, aus 2 kleinen Quadraten bestehend. Auf den Seiten zahlreiche helle Flecken rhombischer oder ovaler Form in ein Netzwerk von schwarzen Linien gefasst. Auf der Schwanzoberfläche 3 grosse braune Rautenflecke. Auge mit schwarzen kräftigen Radien. Von der vordern Lippengegend bis zur Schulter eine breite orangefarbene Zone, oben durch einem kräftigen schwarzen Strich gesäumt. Unterseite blass, gelbgrün.


Calotes mystaceus var. ceylonensis (to new species?) (Plate 3) Kumbukan-aar (South East Ceylon) and North East Province Ceylon from the Messrs. Sarasin. [2] (See Plate III.) The excellently preserved specimens at hand show several not insignificant deviations from the existing descriptions. (DB., Gü. R. b. J., Boul. cat.). Scales of the top of the head smooth, formed as tiles, each equipped with a collar of granules along the free edge; scales of the supraorbital region considerably larger; above the tympanum on each side bearing 2 spines at distance. Tympanum at least ½ the diameter of the orbit. – No gular sack. – Throat scales very strongly carinated, much smaller than dorsal scales (8:5). Four to five feeble spines on the neck; dorsal ridge without any crest or denticulation. Around the middle of the body 60 scales. Dorsal scales barely carinated, some completely smooth, 1 ½ times as large as the abdominal scales. Up to the middle of the body all scales are directed upwards and backwards, on the rear half almost straight backwards directed. Ventral scales mucronate. The aligned rear limb reaches up to the rear eye border. Fourth and fifth fingers equally long. From the eye to the tympanum a row of slightly larger scales. Greenish; across the back 6 large saddle-like cross-blotches, of which the 3 anterior orange coloured follow immediately after each other and are delimited by dark lines, the 3 posterior more spaced, indistinct and weakly coloured. Also across the head runs several light bands. At the beginning of the neck ridge a black double spot composed of 2 small squares. On the sides numerous bright spots of rhombic or oval form, contained in a network of black lines. On the upper side of the tail 3 large brown diamond-shaped spots. Eye with strong black radii. From the anterior lip area to the shoulder a broad orange coloured zone, above margined with a thick black line. Underside pale, yellowish green.

Otocryptis wiegmanni Wagler, 1830 (Plate: 3) Wagler, J., Natürliches System der Amphibien, mit vorangehender Classification der Säugethiere und Vögel., 1830, 150.

Material examined: No type material was designated by Wagler, 1830 (Examined the holotype of Otocryptis bivittata) Male (60.0 mm SVL); Cat. no. ZMB 708; Loc. Ceylon (=Sri Lanka); Coll. Bloch; Date. Unknown (Fig. 17, 18)


OTOCRYPTIS 3) Wiegm., Schluſsohr. Nares Ophryoessae; aures latentes; vertex squamis subcarinatis; gula laevis; digiti fimbriati; cauda teres. (America.) Species: Otocryptis Wiegmanni 4) mihi. 3) Ούς auris, et κρυπτος occultus 4) Gewiſs nicht Azaras Caméléon second.


OTOCRYPTIS 3) Wiegm., Hiddenear. Nostrils like Ophryoessa; ears concealed; scale tops subcarinate; throat smooth; fingers threadlike; tail rounded. (America.) Species: Otocryptis Wiegmanni 4) mine 3) Ούς ear, and κρυπτος hidden 4) certainly not Azara’s second Chameleon.

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Ceratophora stoddartii Gray, 1835 (Plate: 3) Gray, J. E., Illustrations of Indian Zoology, II, 1835: pl.68, fig. 2. (Fig. 19)

Material examined: Holotype Male (67.8 mm SVL); Cat. no. BMNH 1946.8.27.37; Loc. Ceylon (=Sri Lanka); Coll. Stoddart; Date. Unknown. (Fig. 20, 21)

Ceratophora tennentii Günther, 1861 (Plate: 3) Günther, A., in Tennent, J. E., Sketches of the Natural History of Ceylon, 1861: 281, fig. (Fig. 22)

Material examined: One specimen of seven syntypes Male (88.5 mm SVL); Cat. no. BMNH 1946.8.27.32-33; Loc. Ceylon (=Sri Lanka); Coll. Cuming; Date. Unknown. (Fig. 23, 24)

Original description: The specimen in the British Museum is apparently an adult male, ten inches long, and is, with regard to the distribution of the scales and the form of the head, very similar to C. Stoddartii. The posterior angles of the orbit are not projecting, but there is a small tubercle behind them; and a pair of somewhat larger tubercles on the neck. The gular sac is absent. There are five longitudinal quadrangular, imbricate scales on each side of the throat; and the sides of the body present a nearly horizontal series of similar scales. The scales on the median line of the back scarcely form a crest; it is, however, more distinct on the nape of the neck. The scales on the belly, on the extremities, and on the tail are slightly keeled. Tail nearly round. This species is more uniformly coloured than C. stoddartii; it is greenish, darker on the sides.

Ceratophora aspera Günther, 1864 (Plate: 4) Günther, A., The Reptiles of British India, 1864: 131, pl. XIII, fig. G, G’. (Fig. 25)

Material examined: One of four syntypes Male (28.5 mm SVL); Cat. no. BMNH 1946.8.30.51-52; Loc. Ceylon (=Sri Lanka); Coll. Cuming; Date. Unknown. (Fig. 26, 27)

Original description: Head covered with very small, irregular shields, each of which is elevated into a small tubercle; a larger tubercle behind the supraciliary edge, and another on each side of the occiput; occiput with a pair of low ridges, convergent interiorly. Nasal appendage cylindrical, slender, covered with small, imbricate, strongly keeled scales; it is nearly half as long as the head in the male, but quite rudimentary in the female. Labial shields numerous; throat with small, strongly keeled scales, without appendage; no fold in front of the shoulder. Nostril small, lateral. Scales on the back and sides very small, with numerous, irregularly scattered, larger keeled scales; no crest whatever, but some of the larger scales form short angular series across the vertebral line, with their angles pointing backwards. Ventral scales strongly keeled; preanal region covered with very small scales. Tail of moderate length, not compressed, with all its scales keeled; those at its lower surface are scarcely longer than broad. Limbs rather long, the hind limbs extending to, or nearly to, the orbit, if laid forwards. Brownish, marbled with darker; a rhombic light-coloured spot on the sacral region. The brown spots on the fore leg are edged with white in the male. I have examined a male and female of this extraordinary species, both apparently mature, but not longer than 3 inches, of which the tail measures one-half. The British museum received them from Ceylon, from the same source as the C. stoddartii and C. tennentii; hence it is probable that it is also confined to the mountain parts of the interior of the island.

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Figure G of Plate XIII, represents the female in a position which we have observed in many Agames; figure G’ the head of the male.

Lyriocephalus scutatus (Linnaeus, 1758) (Plate: 4) Linnaeus, C., Systema naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tomus I. Editio decima, reformata. 201 Material examined: Iconotype Seb. Mus. vol. I. pl. 109. fig. 3. Loc. Asia; Date. Unknown (Fig. 28)

Original description: ſcutata. 4. L. cauda ſubcompreſſa mediocri, ſutura dorſali dentata, occipite bimucronato. Seb. muſ. I. p. 173. t. 109. f. 3. Salamandra prodigioſa amboinenſis ſcutata. Habitat in Aſia.


Lizard with tail moderately compressed, dorsal suture denticulate, occiput bimucronate [with two points] Seba’s Museum. [Vol.] I. page 173. pl.. 109. fig. 3. Prodigious shielded lizard from Ambon Habitat in Asia.

Cophotis ceylanica Peters, 1861 (Plate: 4) Peters, W. C. H., Monatsberichte der Königlichen Akademie der Wissenschaften zu Berlin, 1861: 1103-1105. Material examined: Lectotype (see Marx, 1958) Male (63.0 mm SVL); Cat. no. ZMB 4240; Loc. Ceylon (=Sri Lanka); Coll. Nietner; Date. Unknown. (Fig. 29, 30)


COPHOTIS nov. gen. 1) Diese Gattung der Iguanoiden schlieſst sich durch ihre zusammengedrückte Körpergestalt, durch die gleiche Zahl der Finger und Zehen und durch das versteckte Trommelfell den Ceylonesischen Gattungen Otocryptis, Lyriocephalus und Ceratophora an. Sie unterscheidet sich von ihnen leicht durch die äuſserst feine Granulation und ungekielte Beschaffenheit der Hand- und Fuſssohle, durch die fast gleiche Länge der 3ten und 4ten Zehe, durch den Schuppenkamm, welcher sich über den ganzen Rücken hinzieht, durch die groſsen Schuppen des Schwanzes und durch die gestrecktere Gestalt der Schnauze. Mit Ceratophora stimmt sie am meisten durch die gröſseren Schuppen an den Seiten der Kehle, des Halses und den Körperseiten, mit Lyriocephalus durch die Occipitaldornen und einem kleinen Postoculardorn überein. Cophotis Ceylanica n. sp. Der Kopf ist pyramidal, doppelt so lang wie breit und hoch, die Schnauze an Länge gleich der Distanz der Augen von einander. Die runden Nasenlöcher öffnen sich seitlich in einfachen Schildchen, welche unmittelbar mit den Supralabialia in Verbindung stehen, von dem Rostralschilde durch ein oder zwei, von einander durch drei Reihen convexer Schuppen getrennt werden. Regelmäſsige, aber an Gröſse den auf der Schnauze befindlichen ziemlich gleiche, gekielte Schuppen bilden jederseits einem Supraorbitalbogen, welcher aus sieben Schuppen besteht und sich dem postorbitalen Dorn anschlieſst. Die Supraorbitalbögen werden nur durch eine Reihe Schuppen von einander getrennt und der Bogen wird bis zum Supraorbitalrande durch drei bis vier Reihen Schuppen ausgefüllt, welche von innen nach auſsen an Gröſse abnehmen. Es finden sich jederseits neun Supralabialia, an welche sich eine zweite Reihe kaum kleinerer Schildchen anschlieſst. Die Schuppen der Schläfengegend sind eben so groſs wie die des Vorderkopfes und einige etwas gröſsere zeigen eben so wie diese in der Mitte eine kurze Spitze. Das Hinterhaupt endigt mit zwei kurzen, wie bei Lyriocephalus convergirenden Dornen. Die Augenlider sind ganz von kleinen körnigen Schüppchen bedeckt, nur an den Augenlidrändern erscheinen sie glatter, und auf dem obern Augenlide sicht

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man eine Reihe von drei bis fünf etwas gröſseren platten Schüppchen. Die Submental- und Kehlgegend wird von schwach gekielten Schuppen bedeckt, welche in, den Infralabialia parallelen, Reihen liegen und an Gröſse nach der Mittellinie allmählig abnehmen. Oben zählt man einem kurzen einspitzigen Mittelzahn, auf welchen jederseits zwei eben so kurze, dann ein längerer und dann dreizehn dreispitzige Backzähne folgen; im Unterkiefer fehlt der Mittelzahn und es folgen jederseits auf zwei einspitzige vierzehn dreispitzige Zähne. Hals, Rumpf und Schwanz sind comprimirt und letzterer bei beiden vorliegenden Exemplaren nach unten gebogen (Greifschwanz?). Der Rücken, die Seiten des Halses, des Rumpfes und der Schwanz sind mit groſsen dachziegelförmig liegenden Schuppen bedeckt, welche in der Mitte der Körperseiten besonders groſs sind; an der Unterseite des Halses, wo sich bei dem kleinen Exemplar ein unbedeutender Kehlsack findet, und an der Brust sind die Schuppen am kleinsten, aber eben so wie die etwa um die Hälfte gröſseren Bauchschuppen gekielt. Auf dem Nacken bilden drei bis vier spitze lange Schuppen einem Kamm, welcher sich durch ähnliche, aber einzeln stehende Schuppen bis zur Kreuzgegend fortsetzt. Der Schwanz hat nichts von einem dorsalen Kamm, aber zwei untere Kiele wie Ceratophora. Die Extremitäten, welche kürzer als bei den verwandten Gattungen erscheinen, sind auf ihrer oberen und äuſseren Seite mit groſsen Schuppen bedeckt, zeichnen sich aber sehr von denen der verwandten Gattungen dadurch aus, daſs die Schuppen der Hand- und Fuſssohlen äuſserst klein sind und die Sohlen der Finger und Zehen gar nicht oder nur sehr schwach, der geringen Gröſse der Schuppen entsprechend, gekielt sind. – Die Farbe ist braun (an den entschuppten Stellen blau); von der Schnauzenspitze längs dem Rande der Oberlippe geht eine gelbliche Binde bis zur Schulter, wo sie plötzlich breiter wird; ein länglicher Fleck hinter jedem Auge, ein gröſserer Nackenfleck vor dem Nackenkamm, ein groſser dreieckiger nach dem Rücken hin spitzer Seitenfleck gleich hinter der vorderen Extremität und breite etwas undeutliche Querbinden am Schwanze sind ebenfalls gelb. Auch die Kehle ist gelb, aber jederseits durch quer von dem Unterlippenrande ausgehende, unregelmäſsige Binden ausgezeichnet. Totallänge 0m, 136; Kopf 0m, 018; Schwanz 0m, 075; vordere Extremität 0m, 023; hintere Extremität 0m, 027; Breite des Kopfs 0m, 008. Die beiden Exemplare sind von Hrn. Nietner auf Ceylon gesammelt worden.

English translation: COPHOTIS nov. gen. 1) This genus of the iguanoids is allied to the Ceylonese genera Otocryptis, Lyriocephalus and Ceratophora for reason of the compressed body, by the same number of fingers and toes, and by the hidden tympanum. It is easily distingushed from them by the extremely fine granulation and the non-carinated condition of the hand and foot soles, by the almost identical length of the 3rd and 4th toe, by the scale ridge which runs along the whole back, by the large scales of the tail, and by the more elongated shape of the snout. It agrees best with Ceratophora by the larger scales on the side of the throat, the neck and the body sides, with Lyriocephalus by the occipital spines and a small postocular spine. Cophotis Ceylanica n. sp. The head is pyramidal, twice as long as wide and high, the snout length identical to the distance between the eyes. The round nostrils open up to the side into simple small shields which are immediately connected to the supralabials, separated from the rostral shield by one or two convex scales, separated from each other by three rows of convex scales. Regular, but in size quite similar to those present on the snout, carinated scales form on each side a supraorbital arch which consists of seven scales and which connects to the postorbital spine. The supraorbital arches are only separated by one row of scales and the arch up to the supraorbital margin is filled by two to three rows of scales which decline in size from inside to outside. On each side there are nine supralabials, to which connects a second row of barely smaller shields. The temple scales are as large as those of the front ones, and some of the larger ones also show a small point in the middle. The occiput ends with two short converging spines as in Lyriocephalus. The eyelids are totally covered by small grainy scales, only at the eyelid margins they appear to be smoother, and on the upper eyelid one can see a row of three to five somewhat larger flat scales. The submental and throat area is covered by weakly carinated scales which lie in parallel rows to the infralabials and are gradually reduced in size after the midline. In the upper jaw one counts one short, unicuspid median tooth, followed on each side by two equally short ones, then a longer one and then thirteen tricuspid molar teeth; in the lower jaw the median tooth is absent and on each side follows after two unicuspid teeth fourteen tricuspid molar teeth. Neck, trunk and tail are compressed, and the latter is bent downwards in the two specimens at hand (grasping tail?). The back, the side of the neck, the trunk, and the tail are covered by large scales arranged like roof tiles, and which are particularly large on the middle of the body sides; on the lower side of the neck, where an

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insignificant throat pouch is found in the small specimen, and on the chest the scales are the smallest, but even so carinated just as the abdominal scales half the size larger. On the neck three to four large pointed scales form a ridge, which continues with similar, but isolated scales up to the small of the back. The tail bears no dorsal ridge but two lower ridges as in Ceratophora. The limbs, which seem to be shorter compared to the related genera, are covered by large scales on the upper and outer sides, but they set themselves apart against the related genera by the extremely small scales of the hand and foot sole and that the soles of the fingers and toes show no or only very weak carination in accordance with the small scale size. – The colour is brown (blue where scales removed); from the snout tip along the margin of the upper lip runs a yellowish band up to the shoulder, where it widens abruptly; an elongate spot behind each eye, a larger nape spot in front of the neck ridge, immediately behind the anterior limb a larger triangular lateral blotch which more pointed toward the back, and broad somewhat indistinct crossbands on the tail, are also yellow. The throat is also yellow, but on each side marked by irregular bands originating from the margin of the lower lip. Total length 0m, 136; Head 0m, 018; Tail 0m, 075; front limb 0m, 023; rear limb 0m, 027; head width 0m, 008. Both specimens were collected by Mr. Nietner on Ceylon.

Sitana ponticeriana Cuvier, 1829 (Plate: 4) Cuvier, Règne Animal, 2nd ed. ii, 1829: 43. Material examined: Holotype Male (46.4 mm SVL); Cat. no. MNHN 6901; Loc. Pondicherry - India; Coll. Leschenault; Date. Unknown (Fig. 31, 32).


LES SITANES. (SITANA. CUV.) (2) Ont, comme les dragons, des dents d’agames et quatre canines; le corps et les membres couverts d’écailles imbriquées et carénées; les cuisses sans pores; mais leurs côtes ne s’étendent point. Ils se distinguent par un énorme fanon qui se porte jusque sous le milieu du ventre, et a plus du double de la hauteur de l’animal. L’espèce connue (Sit. Ponticeriana. Cuv.) est petite, fauve, et a le long du dos une série de grandes taches rhomboidales brunes. Elle vit aux Indes orientales. C’est peut-être de cette tribu des Agamiens que l’on doit rapprocher un reptile fort extraordinaire, qui ne se trouve plus que parmi les fossils d’anciennes couches jurassiques.

English translation: They have, like the dragons, teeth of agamas and four canines: the body and limbs are covered with imbricate and keeled scales; the thighs without pores; but their sides are pointed. They are distinguished by an enormous dewlap which reaches to almost below the middle of the belly, and is more than double the height of the animal. The known species (Sit. Ponticeriana Cuv.) is small, tan, and along the back a series of large brown rhomboidal blotches. It lives in East India. Perhaps it is with this tribe of agamas that one must reconcile a very special reptile, which is no longer found except among the fossils of ancient Jurassic layers. Acknowledgements Several people have directly or indirectly helped to enrich this document. We are very grateful to express our sincere thank to Richard Wahlgren, Aaron M. Bauer, Enrique La Marca, Indraneil Das, Mohomed M. Bahir and anonymous reviewer for reviewing the manuscript and providing valuable comments to improve the document. We are deeply grateful to Mark Oliver Rödel (ZMB) and Frank Tillack (Berlin) who helped to locate much of the .

Literature Cited Andersson, L. G. 1900. Catalogue of Linnean type-specimens of Linnæus's Reptilia in The Royal Museum in Stockholm. Bihang till Kongl. Svenska Vetenskaps-Akademiens Handlingar, 26, 4 (1): 1-29.

Bahir, M. M. and K. P. Maduwage, 2005. Calotes desilvai, a new species of agamid lizard from older literature for us and helped in many other

ways. Also we offer a special word of thank to Bodil Kajrup (NRM), Mark-Oliver Rödel and Rainer Günther for kindly taking photographs of types.

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Morningside Forest, Sri Lanka. The Raffles Bulletin of Zoology, Supplement, 12: 381-392. Bahir, M. M. and A. Silva, 2005. Otocryptis nigristigma, A new species of Agamid Lizard from Sri Lanka. The Raffles Bulletin of Zoology, Supplement, 12: 393-406. Bahir, M. M. and T. D. Surasinghe, 2005. A conservation assessment of the agamid lizards of Sri Lanka. The Raffles Bulletin of Zoology, Supplement, 12: 407-412. Bossuyt, F., M. Meegaskumbura, N. Beenaerts, D. J. Gower, R. Pethiyagoda, K. Roelants, A. Mannaert, M. Wilkinson, M. M. Bahir, K. Manamendra-arachchi, P. K. L. Ng, C. J. Schneider, O. V. Oommen and M. C. Milinkovitch, 2004. Local endemism within the Western Ghats – Sri Lanka Biodiversity Hotspot. Science, 306: 479-481.

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Boulenger, G. A., 1885. Catalogue of the Lizards in the British Museum, Second edition, I, London: xii+436.

Brygoo, E. R., 1988. Les types d’Agamidés (Reptiles, Sauriens) du Muséum national d’Histoire naturelle. Catalogue critique. Bulletin du Muséum national d’Histoire naturelle, 4e Sér. (A), 10 (3), supplement: 1-56.

Cuvier, G., 1829. Le Régne animal distribué d'après son organisation, pour servir de base a l'histoire naturelle des animaux et d'introduction a l'anatomie comparée. Nouvelle édition, revue et augmentée, Tome II. Déterville & Crochard, Paris: xv+406.

Daudin, F. M., 1802. Histoire naturelle, génerale et particulière des Reptiles, vol. III. F. Dufart, Paris: 452.

Duméril, A. M. C. and G. Bibron, 1837. Erpétologie générale our Histoire naturelle complète des reptiles. Tome quatrième. Libraire encyclopedique de Roret, Paris: 571+1. Duméril, M. C. and A. Duméril, 1851. Caralogue méthodique de la collection des reptiles. Gide et Baudry, Paris: 224.

de Silva, A., 2006. Current status of the Reptiles of Sri Lanka. In: Bambaradeniya, C. N. B. (Ed.). Fauna of Sri Lanka: Status of Taxonomy, Research and Conservation. the World Conservation Union, Colombo, Sri Lanka and Government of Sri Lanka: 134-163. Gray, J. E., 1833-1835. Illustrations of Indian Zoology; chiefly selected from the collection of Major-General Hardwicke, F.R.S., Vol. II. Adolphus Richter and Co., London: 263. Günther, A. 1861. in Tennent, J. E., Sketches of the Natural History of Ceylon with narratives and anecdotes illustrative of the habits and instincts of the Mammalia, Birds, Reptiles, Fishes, Insects, &c. including a Monograph of the Elephant and a description of the modes of capturing and training it. Longman, Green, Longman, and Roberts, London: xiii+500. Günther, A. 1864. The Reptiles of British India. The Ray Society, London: xxvii+452. Günther, A., 1872. Descriptions of some Ceylonese Reptiles and Batrachians. Annals and Magazine of Natural History, 4 (9): 85-88. Karunarathna, D. M. S. S., A. A. T. Amarasinghe and E. Stöckli, 2009. Taxonomical, biological and ecological study on Calotes ceylonensis Müller, 1887 (Reptilia: Agamidae) of Sri Lanka. Bonner Zoologische Beiträge: (in press). Kuhl, H., 1820. Beiträge zur Zoologie und vergleichenden Anatomie. Hermann, Frankfurt am Main, Germany. Part I & II: 152. Lescure, J. and C. Marty, 2000. Atlas des Amphibiens de Guyane. MNHN Paris, Collection Patrimoines Naturels, 45: 388. Linnaeus, C., 1748. Systema naturæ sistens regna tria naturæ, in classes et ordines genera et species redacta tabulisque ænis illustrata. Editio sexta. 6th ed. Impensis Godofr. Kiesewetteri. Stockholmiæ. (4), 224 (27), pls 8. Linnaeus, C., 1749. Amoenitates Academicae seu dissertationes variae physicae, medicae botanicae, [1] Holmiae et Lipsiae: 4+563. Linnaeus, C., 1754. Hans Maj:ts Adolf Frideriks vår allernådigste konungs naturalie samling innehållande sällsynte och främmande djur, som bevaras på kongl. lust-slottet Ulriksdahl beskrefne och afritade samt på nådig befallning utgifne af Carl Linnaeus. Stockholm, XXX+96 + 7.

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Linnaeus, C., 1758. Systema naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tomus I. Editio decima, reformata. Holmiae: 823+1. Lönnberg, E., 1896. Linnean type-specimens of birds, reptiles, batrachians and fishes in the Zoological Museum of the R. University in Uppsala. Bihang till Kongl. Svenska Vetenskaps-Akademiens Handlingar, 22, 4 (1): 1-45. Macey, J. R., J. A. Schulte, II, A. Larson, N. B. Ananjeva, Y. Wang, R. Pethiyagoda, N. Rasteger-Pouyani and T. J. Papenfuss, 2000. Evaluating Trans-Tethys migration: an example using Acrodont lizard phylogenetics. Systematic Biology, 49 (2): 233-256. Manamendra-Arachchi, K., A. de Silva and T. Amarasinghe, 2006. Description of a second species of Cophotis (Reptilia: Agamidae) from the highlands of Sri Lanka. Lyriocephalus, 06, Supplement, 1: 1–8. Meegaskumbura, M., F. Bossuyt, R. Pethiyagoda, K. Manamendra-Arachchi, M. Bahir, M. Milinkovitch and C. Schneider, 2002. Sri Lanka: an amphibian hotspot. Science, 298: 379. Müller, F., 1887. Fünfter Nachtrag zum Katalog der herpetologischen Sammlung des Basler Museums. Verhandlungen der Naturforschenden Gesellschaft in Basel, 8 (2): 249-296. Myers, N., R. A. Mittermeier, G. A. B. da Fonseca and J. Kent, 2000. Biodiversity hotspots for conservation priorities. Nature, 403: 853–857. Peters, W. C. H., 1860. Über einige interessante Amphibien, welche von dem durch seine zoologischen Schriften rühmlichst bekannten österreichischen Naturforscher Professor Schmarda während seiner auf mehrere Welttheile ausgedehnten, besonders auf wirbellose Thiere gerichteten, naturwissenschaftlichen Reise, mit deren Veröffentlichung Hr. Schmarda gegenwärtig in Berlin beschäftigt ist, auf der Insel Ceylon gesammelt wurden. Monatsberichte der Königlichen Akademie der Wissenschaften zu Berlin, (April): 182-186. Peters, W. C. H., 1861. Eine neue Gattung von Eidechsen, Cophotis ceylanica, aus Ceylon. Monatsberichte der Königlichen Akademie der Wissenschaften zu Berlin, (December): 1103-1105. Pethiyagoda, R. and Manamendra-Arachchi, K., 1998. A revision of the endemic Sri Lankan agamid lizard genus Ceratophora Gray, 1835, with description of two new species. Journal of South Asian Natural History, 3: 1-50.

Seba, A., 1734. Locupletissimi rerum naturalium thesauri accurata descriptio, et iconibus artificiosissimis expressio, per universam physices historiam. Opus, cui, in hoc rerum genere, nullum par exstitit. Ex toto terrarum orbe collegit, digessit, et depingendum curavit. Tomus I. Wetstenium, Smith & Janssonio-Waesbergios, Amstelaedami: 32+178. Seba, A. 2006. Albertus Seba. Cabinet of Natural Curiosities. The complete plates in colour 1734-1765. New York, Taschen: 588. Tiedemann, F., Häupl, M., Grillitsch, H., 1994. Katalog der Typen der Herpetologischen Sammlung nach dem Stand vom Jänner 1994, Teil II: Reptilia. Kataloge der wissenschaftlichen Sammlungen des Naturhistorischen Museums in Wien, Wien (Naturhistorisches Museum Wien), 10 (Vertebrata 4): 110. Wagler, J. G., 1830. Natürliches System der Amphibien mit vorangehender Classification der Säugethiere und Vögel ein Beitrag zur vergleichenden Zoologie: Mit vorangehender Classification der Säugethiere und Vögel: ein Beitrag zur vergleichenden Zoologie. J. G. Cotta, München, Stuttgart and Tübingen: 354. Wallin, L. 2001. Catalogue of type specimens. 4. Linnaean specimens. Revised version 6. Uppsala. Uppsala University Museum of Evolution Zoology section: 128. Wermuth, H., 1967. Liste der rezenten Amphibien und Reptilien. Agamidae. Das Tierreich, 86. Walter de Gruyter, Berlin: xiv+127. Wiegmann, A. F. A., 1831. Otocryptis. Isis, 24: columns 293-294. Zug, G. R., H. H. K. Brown, J. A. Schulte and J. V. Vindum, 2006. Systematics of the Garden Lizards, Calotes versicolor group (Reptilia: Squamata: Agamidae), in Myanmar: central dry zone population. Proceedings of the California Academy of Sciences, Fourth Series, 57 (1): 1-33.

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MUTUALISM IN Ramanella nagaoi MANAMENDRA-ARACHCHI &

PETHIYAGODA, 2001 (AMPHIBIA: MICROHYLIDAE) AND Poecilotheria

SPECIES (ARACNIDA: THEREPOSIDAE) FROM SRI LANKA

Submitted: 04 November 2008, Accepted: 05 January 2009

D. M. S. Suranjan Karunarathna 1 and A. A. Thasun Amarasinghe

2,3

1 IUCN – Sri Lanka Country office, No: 53, Horton place, Colombo 07, Sri Lanka

2 Taprobanica Nature Conservation Society, No: 146, Kendalanda, Homagama, Sri Lanka

3 Corresponding author: [email protected]

Abstract Ramanella nagaoi is an endemic and vulnerable species of Family Microhylidae distributed in lowland wet

zone rain forests and reproduce and live in and around tree hollows. This article presents some aspects of

mutualism of R. nagaoi with Poecilotheria ornata and P. (cf.) subfusca thereposid spiders in random field visits during the past seven years. This is the first observation of this symbiotic behaviour.

Key words: Ramanella nagaoi, Poecilotheria, commensalism, inter-specific association, symbiotic relationship, Sri Lanka

Introduction The frog Family Microhylidae in Sri Lanka

represents four genera and ten species, while the

genus Ramanella represents four species; R.

variegate (Stoliczka, 1872), R. obscura (Günther,

1864), R. palmata Parker, 1943 and R. nagaoi Manamendra-Arachchi & Pethiyagoda, 2001

respectively. R. palmata, R. obscura and R. nagaoi

are endemic, and R. nagaoi is evidently restricted to

Southern Sri Lanka (Manamendra-Arachchi &

Pethiyagoda, 2001) and it is known from few other

forests (e.g. Sinharaja, Kithulgala, Athwelthota,

Gilimale, Kottawa-Kombala Forest near Hiyare,

Nakiyadeniya, Dediyagala, Hiniduma, Beraliya-

Elpitiya and low altitudes in Sri Pada Nature

Reserve). Here we describe observations of

mutualism relationship between a microhylid frog,

TAPROBANICA, ISSN 1800-427X. April, 2009. Vol. 01, No. 01: pp. 16-18.


tree hollows in the Kanneliya Forest Reserve in Ramanella nagaoi and two species of tarantula

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MUTUALISM IN Ramanella nagaoi AND Poecilotheria SPECIES FROM SRI LANKA


spiders; Poecilotheria ornata and P. (cf.) subfusca.

Previous studies implicate few observations on

commensalism interaction between microhylid

frogs and spiders and the use of chemical cues used

by the spider to recognize the frog (Crocroft & Hambler, 1989; Scakany, 2002). According to

Siliwal & Ravichandran (2008) they observed the

microhylid frog Kaloula taprobanica and the mygalomorph spider Poecilotheria

hanumavilasumica sharing a tree hole in a tamarind

tree in a private plantation on Rameshwaram Island.

Miller (2003) examined another microhylid frog,

Hamptophryne boliviana witnessed communing

with the theraphosid spider Xenesthis immanis.

Scakany (2002) observed the microhylid frog

Chiasmocleis ventrimaculata and its burrow mate, a

theraphosid spider. However, all the previous

studies explain this relationship as a commensalism and here we show this relationship is more

advanced and suggest it may be mutualism, instead.

Commensalism is a close association between two

living organisms of different species, which is

beneficial to one (the commensal) and does not

affect the other (the host) (Taylor et al., 1997).

There are few examples of commensalism: some

orchids or ferns on trees, egret and cow, shark and

remora, Clown fish and sea anemone (Atwaroo-Ali,

2003; Siliwal & Ravichandran, 2008; Taylor et al., 1997). Commensalism (com= together; mensa=

table) means literally “eating at the same table” and

is used to describe symbiotic relationships which do not fit conveniently in to the mutualism and

parasitism categories (Taylor et al., 1997).

However, according to this observation there is a

close association between two living organisms (R.

nagaoi and Poecilotheria species) of different

species, which is beneficial to both parties.

Observations We recorded the mutualistic association between R.

nagaoi and Poecilotheria species from

Bambarabotuwa-Ratnapura (alt. 950 m; 6o 39’

53.35 N, 80o 35’ 01.54 E), Kithulgala (alt. 120 m; 7o

00’ 00.20 N, 80o 24’ 59.86 E), Pompekale-

Ratnapura (alt. 85 m; 6o 41’ 04.70 N, 80

o 24’ 20.72

E), Athwelthota (alt. 210 m; 6o 32’ 17.48 N, 80o 17’

03.49 E), Gilimale (alt. 390 m; 6o 45’ 35.92 N, 80

o

27’ 13.51 E), Kottawa-Kombala near Hiyare (alt. 70

m; 6o 05’ 44.48 N, 80o 18’ 38.04 E), Kanneliya (alt.

200 m; 6o 13’ 36.87 N, 80o 24’ 05.17 E), Hiyare

(alt. 120 m; 6o 03’ 37.15 N, 80

o 19’ 27.93 E),

Nakiyadeniya (alt. 180 m; 6o 11’ 30.38 N, 80o 22’

39.89 E), Dediyagala (alt. 240 m; 6o 11’ 16.22 N,

80o 23’ 59.68 E), Hiniduma (alt. 260 m; 6

o 20’

450 m; 6o 25’ 11.61 N, 80o 27’ 05.43 E), Beraliya-

Elpitiya (alt. 150 m; 6o 15’ 48.58 N, 80o 12’ 22.39

E) and low altitudes in Sri Pada Nature Reserve (alt.

400 m; 6o 46’ 56.44 N, 80o 27’ 26.11 E), all in Sri

Lanka, in random field visits during the past seven

years.

During these surveys we located 17 (17.35%) tree

holes containing both of these species out of 98 tree

holes where only either Poecilotheria species or R.

nagaoi inhabited. During these surveys we located

66 tree holes with R. nagaoi (alone or with

Poecilotheria species). Out of them 32 tree holes

presence of eggs or tadpoles of R. nagaoi.

Additionally, 34 tree holes were recorded with

Poecilotheria species (alone or with R. nagaoi). Out

of them, 17 tree holes had presence of eggs or

juveniles of Poecilotheria species. During this

survey we observed 12 tree holes with eggs and

tadpoles or juveniles of both species in one tree

hole, out of 17 tree holes where both species live

together.

The natural predators on eggs of Poecilotheria

species are mantids, ants and other spider species;

for juveniles: mantids, Hemidactylus depresses,

Boiga species and birds; for adults: Boiga species

and birds. Natural predators of R. nagaoi eggs are Hemidactylus depresses, ants and Boiga species (De

Silva (2006) observed Cercaspis carinatus occupied

in the habitat of R. nagaoi); for juveniles: Hemidactylus depresses and for adults: birds.

However we didn’t observe any mantid, ant, other

spider species or Hemidactylus depresses and Boiga

juveniles in the tree holes where both R. nagaoi and

Poecilotheria species inhibited during our random

field visits. However, birds and adult Boiga species

were observed while predating. We observed

several times that the Poecilotheria species attacked

on H. depresses while they were trying to eat R. nagaoi eggs (and De Silva (2006) also documented

H. depresses feeds on R. nagaoi eggs. Furthermore,

we also have seen many times R. nagaoi feeding on ants while they were attacking Poecilotheria species

eggs. In addition, we have trace in many occasions

the body parts of the preys fell in to the water while

Poecilotheria species were feeding. Sometimes this

nutrition may help for the survival of R. nagaoi

tadpoles. Accordingly, we believe this relationship

is advanced and it is not limited and depending

strictly upon feeding. In conclusion, we suggest

considering this relationship as mutualism, where

both species gain benefits like food, and protection from predators, among others.

08.13 N, 80o 18’ 20.66 E), Koskulana-Panapola (alt.

17

KARUNARATHNA AND AMARASINGHE, 2009


Acknowledgements The authors wish to thank Enrique La Marca, Tzi

Ming Leong and Mohomed M. Bahir for reviewing

the manuscript. Then the first author would like to express his sincere thank to Mendis Wickramasinghe

for providing valuable comments. Finally we would like to thank Dinesh Gabadage (TNCS) and Niranjan Karunarathna for their help.

Literature Cited Atwaroo-Ali, L., 2003. Macmillan CXC Science Series

Biology. Macmillan Publishers Limited, Malaysia:

100.

Crocroft, R. B. and K. Hambler, 1989. Observations of

a commensal relationship of the microhylid frog

Chiasmocleis ventrimaculata and the burrowing

theraphosid spider Xenesthis immanis in southerastern

Peru. Biotropica, 21 (1): 2-8.

De Silva, M. A., 2006. An experimental phytotelm

conservation project for a Microhylid frog Ramanella

nagaoi. Loris, 24 (3&4): 15-19.

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

Ramanella nagaoi, a new tree-hole frog

(Microhylidae) from southern Sri Lanka. Journal of

South Asian Natural History, 5 (2): 121–133.

Miller, G., 2003. Observations of commensalism

between burrowing theraphosid spiders and the

microhylid frog species, Hamptophrynes boliviana.

CNR Student Research Symposium, College of Natural

Resources, University of Wisconsin - Stevens Point:

24.

Scakany, J., 2002. Study on the Chemical

Communication between the Microhylid Frog,

Chiasmocleis ventrimaculatata, and a Theraphosid

Spider involved in a Commensal Relationship. Masters

dissertation report submitted to State University of

New York: 13.

Siliwal, M. and B. Ravichandran, 2008. Commensalism in Microhylid frogs and Mygalomorph

spiders. Zoos’ Print Magazine, 23 (8): 13.

Taylor, D. J., N. P. O. Green and G. W. Stout, 1997. Biological Science: Soper, R. (Ed.). Cambridge

University Press, UK: 984.

18


ON DISTRESS CALLS OF MALE Hemidactylus brookii parvimaculatus DERANIYAGALA, 1953 (REPTILIA: GEKKONIDAE) FROM SRI LANKA

Submitted: 09 September 2008, Accepted: 15 September 2008

Dieter Gramentz Földerichstraße 7 D-13595 Berlin, Germany: E-mail: [email protected] Abstract Two types of distress calls were recorded from three male Hemidactylus brookiii parvimaculatus. One type consisted of clicks and one of a single squeak. Twelve calls were analysed. Distress calls comprising a squeak had an average length of 0.070 sec while those comprising clicks averaged 0.329 sec. The difference in length of the two types of calls was significant (P<0.05). Average maximum sound intensity of all calls was 76.9 dB and was reached between 3045 Hz and 7473 Hz (x = 4451 Hz). Maximum frequency varied from 7194 Hz to 16238 Hz having an average of 13393 Hz and the average minimum frequency was 1230 Hz. All squeak distress calls showed harmonics, whereas no harmonics occurred in calls with clicks. Key words: Hemidactylus brookii parvimaculatus, bioacoustics, distress call, Sri Lanka. Introduction A number of gecko species are known to emit calls in potentially dangerous situations. These may either be threat calls of a gecko calling prior to physical contact and to intimidate a conspecific or a potential predator. These calls may be rather long in duration. After contact, usually after a gecko has been bitten, it may react with a comparatively short distress call which is certainly the call most often described in geckos (e.g. Barts, 2002, 2006; Brown, 1985; Frankenberg, 1973, 1975, 1978; Gramentz, 2005b,c,d, 2007; Gramentz & Barts, 2004; Kreuzer & Grossmann, 2003; Marcellini, 1974; Morgue, 1913; Nettmann & Rykena, 1985; Scerbak, 1981;

Werner et al., 1978). According to Frankenberg (1975) distress calls have an antipredatory function. Information from a number of species of the genus Hemidactylus is available on sound production during different situations, including H. frenatus (Frenkel, 2006; Marcellini, 1974, 1977; McCann, 1940), H. turcicus (Frankenberg, 1982), H. mabouia (Gramentz, 2003; Regalado, 2003), H. platycephalus (Gramentz, 2005a) and H. angulatus (Gramentz, 2005d). The first anatomical studies on the voice apparatus of the genus were carried out one century ago by Steck (1908) on H. garnotii. Probably the first mention of the voice of


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ON DISTRESS CALLS OF MALE Hemidactylus brookii parvimaculatus FROM SRI LANKA


Hemidactylus brookii and at the same time its distress call was made by Lang (in Schmidt, 1919). He reported that the geckos on being captured emitted a weak sound. Despite the wide distribution of H. brookii no bioacoustic analysis of this type of call has been carried out on this species since. The aim of the present work is to fill this gap by an analysis of the sound properties and structure of this part of the species bioacoustic behaviour. Material and Methods Four adult male H. b. parvimaculatus were caught at Aluthgama (alt. 2 m; 06°27’ N, 79°59’ E), Sri Lanka, in November 2007 for sound recordings of distress calls. Altogether, 12 distress calls were recorded from three specimens. The snout – vent length of these geckos were 4.8 cm (male 1), 4.9 cm (male 2) and 5.4 cm (male 3). All calls were emitted in identical situations while handling the gecko, simulating the attack of a predator. The recording equipment is the same as described by Gramentz (2005a,c). The sound card used was Creative Soundblaster Audigy 2 ZS Platinum Pro with a sample rate of 44100 Hz, 16 bit. Various software was used for sound analysis such as Adobe Audition 1.5, Avisoft-SASLab, Creative WaveStudio and Raven1.2. The areas in the 3D figures below about 300 Hz result from working noises of the recorder. Air temperatures at which the calls were recorded ranged from 27.6 – 27.9 °C. Distance between geckos and microphone during recording distress calls was 5 – 10 cm. The sound of the geckos was recorded either during the night of capture or the following morning. The geckos were released the following night at precisely the same locations they had been first seen prior to capture. Terminology used is the same as in Gramentz (2003 & 2008). Results Of four male H. b. parvimaculatus caught for sound analysis three (75%) produced distress calls. The ultimate intention of the call is probably to be released from the supposed predator’s grip. There is some marked variation in the distress calls of H. b. parvimaculatus. However, beside the differences in call types produced in a distress situation, each call type has a rather uniform and identifiable structure. Calls could be separated into two groups according to their structure such as those formed by a highly condensed number of amplitudes with a squeak-like sound (n = 9) and those containing a number of

clicks with a discernable time gap between them (n = 3). The clicks of the call depicted (Pl. 5: Fig. 1) had time gaps of 0.062, 0.033 and 0.020 sec. The duration of four clicks measured a mere 0.003 sec. In another call with three clicks the gaps between them were 0.130 and 0.073 sec, while the clicks varied in length between 0.003 and 0.007 sec. The five clicks of the call (Pl. 5: Fig. 2 & 3) showed time gaps of 0.152, 0.129, 0.120 and 0.183 sec. The first click appeared 0.055 sec after the beginning of the call. The clicks also had also a short length between 0.004 and 0.006 sec. According to the frequency of recorded distress calls, those with a squeak occur more frequently representing 75% of the calls and the remaining 25% of calls were those with clicks. All calls were rather short in duration, always less than one second and usually less than one quarter second in length. Distress calls without clicks had an average length of only 0.070 sec (SD = 0.03, range: 0.029 – 0.116 sec, n = 9), while those with clicks were considerably longer with an average of 0.329 sec (SD = 0.28, range: 0.121 – 0.642 sec, n = 3). From these data it can also be seen that the longest distress call was one with clicks and a length of 0.642 sec and the shortest was 0.029 sec long with a squeak. There is a statistically significant (P<0.05) difference between the mean length of those calls with a squeak and with clicks (t = 2.57, t- test). All distress calls had an average maximum sound intensity of 76.9 dB. Maximum sound intensity lasted only between 0.002 and 0.004 sec its position was always located in a strong harmonic. Maximum sound intensity was reached between 3045 Hz and 7473 Hz (x = 4451 Hz, SD = 1242, n = 12). Maximum frequency varied from 7194 Hz to 16238 Hz having an average of 13393 Hz (SD = 2861, n = 12). Average minimum frequency was 1230 Hz (SD = 610, range: 277 – 2122 Hz, n = 12). All distress calls with a squeak showed harmonics over most part of the call. No harmonics could be identified in the short clicks. Some of the distress calls show very peculiar harmonics (Pl. 5: Fig. 4 & 5). In particular, one call each of male 2 and male 3 showed peculiar bending or up- and down-curved harmonics. By way of comparison, in other calls (Pl. 5: Fig. 6 & 7)

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harmonics were more or less linear, but kept the rather large frequency interval between harmonics. However, in two distress calls (Pl. 5: Fig. 3 & 10), one with a squeak together with clicks and one without clicks, harmonics were rather linear, more numerous and showed very short intervals between harmonics in comparison to all other distress calls. The call of Pl. 5: fig. 7 was interesting as it contained altogether nine harmonics of three different intensities with 3 harmonics to each category, classifiable as strong, medium and weak. Harmonics of Pl. 5: fig. 7 analysed from bottom to top: the lowest harmonic is of medium strength at about 1568 Hz, followed by three strong harmonics at 3137 Hz, 4797 Hz and 6181 Hz. Another two medium strong harmonics followed at 7658 Hz and 9226 Hz and on top three weak harmonics at 10518 Hz, 12271 Hz and 13747 Hz. The average frequency interval between these harmonics was rather homogeneous being 1522 Hz (SD = 147.8). Average maximum sound intensity (dB) of weak, medium and strong harmonics was 46.3 dB (SD = 2.3, n = 3), 59.3 dB (SD = 3.4, n = 3) and 79.3 dB (SD = 4.4, n = 3) respectively. Interestingly, the frequency of the lowest harmonic is virtually identical to the average frequency interval between the harmonics. The maximum frequency in all recorded distress calls varied between 7194 Hz and 16238 Hz with an average of 13393 Hz (SD = 2861, n = 12) and the average minimum call frequency was 1230 Hz (SD = 610, range: 277 – 2122 Hz, n = 12). Discussion In comparison to the distress calls of other gecko species described in the literature the calls of H. brookii parvimaculatus are also short, but some variations in length and structure exist between species. Very short distress calls have been recorded in Haemodracon riebecki with an average length of only 0.069 sec (range: 0.046-0.080 sec, Gramentz, 2005b) and also Stenodactylus stenurus with an average of 0.034 sec (range: 0.033-0.036 sec, Gramentz, 2004). However, in S. stenurus three types of distress calls were found and the longest had an average length of 0.129 sec (range: 0.111-0.143 sec). In Thecadactylus rapicauda an average distress call length of 0.235 sec was found (range: 0.091-0.360 sec, Gramentz, 2007). In terms of call lengths the distress calls of the closely related Hemidactylus angulatus were also shorter than one second (with few known exceptions distress calls of gecko species studied to date frequently have

lengths of 1/10 to 3/10 of a second), but they are considerably longer (x = 0.454 sec, range: 0.224 – 0.955 sec, Gramentz, 2005d) than those of H. b. parvimaculatus. Furthermore no distress call with clicks could be recorded for H. angulatus. Clicks have been found only in the advertisement call for H. angulatus (Gramentz, 2005d). However, beside some marked differences in distress calls between the related taxa H. b. parvimaculatus and H. angulatus, there are also similarities in the calls of both. While in all distress calls of H. b. parvimaculatus harmonics were present, in H. angulatus this was also the case in 7 of 12 (58%) calls (Gramentz, 2005d). Beside differences in sound intensity and maximum frequency the type of distress call of H. b. parvimaculatus shown in Pl. 5: fig. 8, 9 & 10 can be similar to that of T. rapicauda (Gramentz, 2007), but some calls are different in the respect that amplitudes show different strength over the call length (Pl. 6: Fig. 11). When comparing the structure of the distress calls of H. angulatus with that of H. b. parvimaculatus there are obvious differences, which can be seen in oscillograms. While in H. angulatus a distress call was formed by 2 to 8 pulses (x = 5.8) (Gramentz, 2005d), no pulse structure was evident in the calls of H. b. parvimaculatus. A pulse structure is also present in the distress call of H. platycephalus (Gramentz 2005a). Possibly, there are also differences in the frequency range between both species. In male H. angulatus frequencies between around 100 Hz and 12500 Hz were recorded which is less in terms of minimum frequency as well as maximum frequency than the present findings of H. b. parvimaculatus. Wever (1978) studied the ear structure of one specimen of H. brookii, however no hearing properties were given. In the related species H. angulatus he found that the ear showed excellent sensitivity over the range of 200-1000 Hz. As it was simulated here, a distress call is most likely not only produced when a gecko is being bitten by a conspecific but also during an attack of a potential predator forming a part of the species anti predator behaviour. Presuming that - as in other gecko species distress calls also have an intraspecific meaning - the sensitivity range should be expected to be slightly higher in H. b. parvimaculatus. Furthermore it is very probable that H. b. parvimaculatus is using acoustic communication in other situations than in the present case and is capable of producing some other types of calls.

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ON DISTRESS CALLS OF MALE Hemidactylus brookii parvimaculatus FROM SRI LANKA


Frankenberg (1982) tentatively described different calls in H. turcicus such as escape call, defence call, release call and threat call produced in distress situations. The two different call types with their high variation found in H. b. parvimaculatus and summarised here as distress calls may, too, have different functions similar to H. turcicus. Further studies on the behaviour of the geckos and their associated call types should be carried out which may possibly lead to our better differentiation of the calls of H. b. parvimaculatus. Acknowledgement I would like to thank Olivier S. G. Pauwels (IRSNB), John Rudge and anonymous reviewers for useful comments. Literature Cited Barts, M., 2002. Die Dickfingergeckos des südlichen Afrikas. Teil II. Die Haltung und Vermehrung des Gebänderten Dickfingergeckos, Pachydactylus fasciatus Boulenger, 1888. Sauria, 24 (1): 3-8. Barts, M., 2006. Pachydactylus haackei Haacke’s Dickfingergecko. Sauria, 28 (1): 54. Brown, A. M., 1985. Ultrasound in gecko distress calls (Reptilia: Gekkonidae). Israel Journal of Zoology, 33: 95-101. Frankenberg, E., 1973. Vocalizations of the fan-toed gecko, Ptyodactylus hasselquistii: Israel Journal of Zoology, 22: 205. Frankenberg, E., 1975. Distress calls of gekkonid lizards from Israel and Sinai. Israel Journal of Zoology, 24: 43-53. Frankenberg, E., 1978. Calls of male and female tree geckos, Cyrtodactylus kotschyi. Israel Journal of Zoology, 27: 53-56. Frankenberg, E., 1982. Vocal behaviour of the Mediterranean house gecko Hemidactylus turcicus. Copeia, 1982: 770-775. Frenkel, C., 2006. Hemidactylus frenatus (Squamata: Gekkonidae): call frequency, movement and condition of tail in Costa Rica. Revista de biologia tropical, 54 (4): 1125-1130. Gramentz, D., 2003. Zur Stimme und Rufperiodik von Hemidactylus mabouia (Moreau de Jonnès, 1818). Sauria, 25 (2): 23-28. Gramentz, D., 2004. Der Schreckruf von Stenodactylus petrii Anderson, 1896. Sauria, 26 (4): 13-16.

Gramentz, D., 2005a. Zur intraspezifischen bioakustischen Kommunikation von Hemidactylus platycephalus Peters, 1854 (Reptilia: Sauria: Gekkonidae). Gekkota, 5: 155-154. Gramentz, D., 2005b. Der Schreckruf von Haemodracon riebeckii Peters, 1882 (Reptilia: Sauria: Gekkonidae). Gekkota, 5: 170-178. Gramentz, D., 2005c. Zum Defensivverhalten und Schrecklaut von Geckonia chazaliae Mocquard, 1895. Sauria, 27 (3): 23-27. Gramentz, D., 2005d. Zur intraspezifischen bioakustischen Kommunikation von Hemidactylus brookiii angulatus Hallowell, 1852. Sauria, 27 (4): 41-46. Gramentz, D., 2007. Zum bioakustischen Verhalten männlicher Thecadactylus rapicauda Houttuyn, 1782. Sauria, 29 (3): 13-18. Gramentz, D., 2008. Zum bioakustischen Verhalten von Ptenopus carpi Brain, 1962. Sauria, 30 (1): 43-46. Gramentz, D. and M. Barts, 2004. Der Schrecklaut von Pachydactylus rugosus A. Smith, 1849. Sauria, 26 (1): 23-26. Kreuzer, M. and W. Grossmann, 2003. Beobachtungen an Gekko ulikovskii Darewski & Orlow, 1994 und Gekko grossmanni Günther, 1994 im Terrarium. Sauria, 25 (3): 3-11. Marcellini, D., 1974. Acoustic behavior of the gekkonid lizard, Hemidactylus frenatus. Herpetologica, 30 (1): 44-52. Marcellini, D., 1977. The function of a vocal display of the lizard Hemidactylus frenatus (Sauria: Gekkonidae). Animal Behaviour, 25: 414-417. McCann, C., 1940. A reptile and amphibian miscellany. Journal of the Bombay natural History Society, 41 (4): 742-764. Morgue, M., 1913. Étude sur le Phyllodactylus d’Europe, “Phyllodactylus europaeus” Gené. Bulletin de la Sociéte Linnéenne, Marseille, 1: 45-51. Nettmann, H. K. and S. Rykena, 1985. Verhaltens- und fortpflanzungsbiologische Notizen über kanarische und nordafrikanische Tarentola-Arten. Bonner Zoologische Beiträge, 36 (3&4): 287-305. Regalado, R., 2003. Roles of visual, acoustic, and chemical signals in social interactions of the tropical house gecko (Hemidactylus mabouia). Caribbean Journal Science, 39 (3): 307-320.

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GRAMENTZ, 2009


Scerbak, N., 1981. Cyrtodactylus russowii (Strauch 1887) – Transkaspischer Bogenfingergecko. 75-83. In: H. Böhme (Ed.), Handbuch der Reptilien und Amphibien Europas, Akademische Verlagsgesellschaft, Wiesbaden. Schmidt, K. P., 1919. Contributions to the herpetology of the Belgian Congo based on the collection of the American Museum Congo Expedition, 1909-1915. Part I. Turtles, crocodiles, lizards and chamaeleons. Bulletin of the American Museum Natural History, 39: 385- 624. Steck, L., 1908. Der Stimmapparat des Hemidactylus garnoti Dum. et Bibr. Zoologische Jahrbücher, 25: 611-636. Werner, Y. L., E. Frankenberg & O. Adar, 1978. Further observations on the distinctive vocal repertoire of Ptyodactylus hasselquistii cf. hasselquistii (Reptilia: Gekkoninae). Israel Journal of Zoology, 27: 176-188. Wever, E. G., 1978. The Reptile Ear. Princeton University Press, New Jersey, 1024.

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PRADEEP AND AMARASINGHE, 2009


OVIPOSITIONAL BEHAVIOR OF Calotes ceylonensis MÜLLER, 1887 (REPTILIA: AGAMIDAE) OBSERVED IN THE CENTRAL PROVINCE OF SRI LANKA

Submitted: 10 June 2008, Accepted: 01 July 2008

W. A. A. D. Gayan Pradeep 1 and A. A. Thasun Amarasinghe

1,2

1 Taprobanica Nature Conservation Society, No. 146, Kendalanda, Homagama, Sri Lanka


Abstract This is the first documented observation of the oviposition of Calotes ceylonensis. The ovipositional

behavior consisted of digging of the hole nest to lay the eggs; the laying of the eggs; the scraping of the soil

to bury the eggs; the filling of the spaces between the eggs, and the hole nest; the tight compression of the

soil, and camouflage the nest.

Keywords: Agamidae, Draconinae, Calotes ceylonensis, Egg-laying behaviour, Sri Lanka, Conservation.

Introduction There are eighteen species of agamid lizards in Sri

Lanka, fifteen (83.33%) of them are endemic to the

island (Manamendra-Arachchi et al., 2006). Seven

species out of them belong to the genus Calotes.

Five of them (C. ceylonensis Müller, 1887; C.

liocephalus Günther, 1872; C. liolepis Boulenger,

1885; C. nigrilabris Peters, 1860; C. desilvai Bahir

& Maduwage, 2005) are endemic. The remaining

two, Calotes calotes (Linnaeus, 1758) and C.

versicolor Daudin, 1802 are probably widespread species throughout South East Asia (Taylor, 1953).

According to the published literature, Calotes

ceylonensis is a largely arboreal species found only from the low country dry and intermediate zones

below 500 m a.s.l (Karunarathna et al., 2009). It is rare and a vulnerable species (Bahir & Surasinghe,

2005; Manamendra-Arachchi & Liyanage, 1994;

IUCNSL & MENR, 2007).

The information available on this species is scarce

and, therefore, further studies on their behaviour and ecology are needed, which may be very

important for the conservation of the species



24

OVIPOSITIONAL BEHAVIOR OF Calotes ceylonensis OBSERVED AT THE CENTRAL PROVINCE OF SRI LANKA


(Karunarathna et al., 2009). In this paper we

describe the ovipositional behaviour of C.

ceylonensis, which constitutes the first described

observation of the ovipositioning for this lizard.

Location and Methods of Observation Observations were made approximately 3 km from

Wewala-Dambulla Road in Kaludiya Pokuna near

Kandalama (7o 52.47’ N, 80o 44.08’ E; alt. 228 m

a.s.l.) in Matale District, Central Province, Sri

Lanka. The habitat consisted mainly of disturbed

home gardens. The ground was covered with large

amounts of dry leaf litter and the soil was rough.

There was approximately 80% canopy cover and

clear undergrowth. Observations of the lizard were

made by the naked eye from 2 m away between

10:40 and 12:45 hr. The animal was not disturbed

during observation. All measurements were taken to

the nearest 0.1 mm using digital caliper.

Observations A mature female Calotes ceylonensis (snout to vent

length: 72.1 mm) lying on a Ficus bengalensis tree

(10 m high) 2 m above the ground level, was observed on 08 September 2008 at about 10:40 hr.

The temperature was 28.4 °C and the humidity

53%. The weather was gloomy and the cloud cover

was 7/8.

Digging the hole nest First, the lizard descended down from the Ficus

bengalensis tree and walked away 2 m from the tree

base. Then it was turned its body to make a body pit

(diameter about 12 cm) but after few minutes it

stopped that activity and moved away about 1 m. At

that place it made another body pit as mentioned

above. Again, it gave up that action and moved

another 2 m away. This time it started to make the

body pit near to a Xylopia nigricans tree. It took

approximately 5 minutes to make the body pit,

which was 21 cm away from that tree base. This

lizard used its hind limbs to remove leaf litter while

making the hole. During this activity it pressed and

gripped its lower jaw in addition to the fore limbs.

This grip made easier to remove and throw leaf

litter backwards using both hind limbs fast and

powerfully. After that it lifted the anterior part of its

body using its forelimbs. Then it looked around for

about 5 minutes. During this time it repeatedly

turned its head 180° two times, without moving its

body. During this time it changed its body colour to become darker to match the ground.

continued for approximately 5 minutes. After that it

stopped digging and looked around for

approximately 5 minutes. while repeatedly turning

its head 180° one time, without moving its body.

Again it started the digging and this time the female dug the hole continuously for approximately 20

minutes. It stopped and looked around for about 5

minutes. while turning its head 180° around two

times, without moving its body. After that it

continued to dig the hole for another 20 minutes,

stopping one more time for 5 minutes to rest.

Sometimes it used one forelimb for digging whiles

other forelimb was kept free on the ground. After

short time it changed the fore limbs and dug using

the fore limb that was kept free. The hole was dug

into the ground at a 45° angle. The final hole was

50.8 mm deep and 38.1 mm in diameter.

Laying the eggs After the digging, the female turned its body 180°

clockwise, placing the posterior part of its body

over the hole. It then looked around again. Then it

lifted the posterior part of its body to lay eggs. This

egg laying was very similar to the egg laying of

Calotes liolepis. Three eggs were laid at a rate of one per minute (Fig. 1). The eggs were pure white

and elliptical, with a mean length of 15.5 mm and a

mean width 8.4 mm. After the eggs were laid, the

female lowered its posterior part and looked around.

Then the female packed and placed the eggs below

ground level using the anterior part of its lower jaw.

Fig. 01: The egg laying of C. ceylonensis

Burying the eggs and camouflaging the nest After placing eggs it turned 180° clockwise and

began to drag the soil towards the hole nest using its

fore limbs. The dragged soil was pressed using

anterior half of its lower jaw for half an hour. After

looking around, it dragged the surrounding leaves of

Xylopia nigricans (Family: Annonaceae), Diospyros

The female C. ceylonensis then began digging the

ground while scraping the soil with its forelimbs,

which was thrown backwards beside its body. This

25

PRADEEP AND AMARASINGHE, 2009


ebenum (Family: Ebenaceae), Ficus bengalensis

(Family: Moraceae) and Mesua nagassarium

(Family: Clusiaceae) over the nest site for

camouflage. It remained motionless for 5 minutes

and then ran towards the Ficus bengalensis tree, where we observed it first. Afterwards it was caught

for measurements, and then released.

Discussion The oviposition behavior of this species varies from

the oviposition behavior of Calotes versicolor

(Amarasinghe & Karunarathna, 2007) and C.

liocephalus (Amarasinghe & Karunarathna, 2008).

According to Amarasinghe & Karunarathna (2007),

while laying eggs C. versicolor places its cloacal

aperture over the opening of the nest hole and it

places its hind limbs without expanding them (Fig.

2), but C. ceylonensis places its cloacal aperture

over the opening of the hole and places its hind

limbs expanding and also without much lifting its

posterior part (Fig. 3). According to Amarasinghe &

Karunarathna (2008) C. liocephalus places the

posterior part of the body inside the hole while

laying eggs (Fig. 4). In comparison, the egg layings

of C. versicolor (Amarasinghe & Karunarathna,

2007) and C. liocephalus (Amarasinghe &

Karunarathna, 2008) occurs in wet habitat while this

observation was made in a dry habitat.

Fig. 02: The egg laying of C. versicolor

The C. versicolor lifts the anterior part of the body

with its forelimbs while turning its head to look

around (Amarasinghe & Karunarathna, 2007) and

C. liocephalus coils its entire body inside the hole

while bending the anterior part of its body to look

around (Amarasinghe & Karunarathna, 2008), but

C. ceylonensis only turned head at an angle of 90o to

looking around.

The C. versicolor makes a knocking noise while

packing and placing the eggs in the hole using its

lower jaw (Amarasinghe & Karunarathna, 2007) while the C. liocephalus (Amarasinghe &

Karunarathna, 2008) and C. ceylonensis places them

softly without making any noise.

Fig. 03: The egg laying of C. ceylonensis

C. versicolor and C. liocephalus did not mark the

body pit to dig the hole nest (Amarasinghe &

Karunarathna, 2007, 2008) as C. ceylonensis. C.

ceylonensis threw the soil backward under its body

through its raised hind limbs similar to C. versicolor

and C. liocephalus (Amarasinghe & Karunarathna,

2007).

Fig. 04: The egg laying of C. liocephalus

In this ovipositioning, the female had much effort to

remove leaf litter and it used its lower jaw also to

grip. Additionally, the soil also was comparatively

hard and rough. Therefore it used one forelimb to

dig the soil at a time while keeping other at rest and

then changed them most of times. At that day the

weather was very dry and dull even when there was a rain during the previous day. During this

observation we could trace Otocryptis nigristigma,

Eutropis tammanna and Lankascincus fallax

sympatrically. Nonetheless, the female did not show

any response to these sympatric species while

laying eggs, even when they moved near the hole

nest.

Acknowledgments We wish to thank Enrique La Marca and Indraneil

Das for reviewing the manuscript and Mohomed M.

Bahir for providing valuable comments. The first

author thanks Rajnish Vandercone for his generous

support for the field visits. Finally we would like to

26

OVIPOSITIONAL BEHAVIOR OF Calotes ceylonensis OBSERVED AT THE CENTRAL PROVINCE OF SRI LANKA


thank Madhava Botejue (TNCS) and Dinesh

Gabadage (TNCS) for their help.

Literature Cited Amarasinghe, A. A. T. and D. M. S. S. Karunarathna, 2007. Beobachtungen zum Eiablageverhalten der

Indischen Schönechse Calotes versicolor (Daudin,

1802) (Reptilia: Agamidae) in einem anthropogenen Biotop in Sri Lanka. Sauria, 29 (3): 27–30.

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

2008. Observation on the Oviposition behaviour of the

Crest-less Lizard Calotes liocephalus (Reptilia: Agamida) in the Knuckles forest region of Sri Lanka.

Asiatic Herpetological Research, 11: 13-16.

Bahir, M. M. and T. D. Surasinghe, 2005. A

conservation assessment of the agamid lizards of Sri

Lanka. The Raffles Bulletin of Zoology, Supplement

12: 407-412.

IUCNSL & MENR, 2007. The 2007 Red List of

Threatened Fauna and Flora of Sri Lanka. Colombo,

Sri Lanka. xiii+148.

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

E. Stöckli, 2009. Taxonomical, Biological and

Ecological study on Calotes ceylonensis Müller, 1887

(Reptilia: Agamidae) of Sri Lanka. Bonner Zoologische Beiträge, (in press).

Manamendra-Arachchi, K. and S. Liyanage, 1994.

Conservation and distributions of the agamid lizards

of Sri Lanka with illustrations of the extant species.

Journal of South Asian Natural History, 1 (1): 77-96.

Manamendra-Arachchi, K., A. de Silva and T. Amarasinghe, 2006. Description of a second species

of Cophotis (Reptilia: Agamidae) from the highlands of Sri Lanka. Lyriocephalus, 06, Supplement 1: 1–8.

Taylor, E. H., 1953. A review of the lizards of Ceylon. The University of Kansas science bulletin, 35 (12):

1525-1585.

27

WARAKAGODA AND SIRIVARDANA, 2009


THE AVIFAUNA OF SRI LANKA: AN OVERVIEW OF THE CURRENT STATUS

Submitted: 19 February 2009, Accepted: 31 March 2009

Deepal Warakagoda1 and Udaya Sirivardana1,2

1 Ceylon Bird Club, 127, Nawala Rd, Colombo 5, Sri Lanka 2 Corresponding author: [email protected] Abstract Sri Lanka has a high diversity of avifauna, with 236 breeding and 203 purely migrant species. Among the former, 33 species and a further 68 subspecies are endemic to it. Since the systematic study of the birds of the island began in the 18th century intermittent discoveries of breeding and migrant taxa have led to a gradual increase in the total number of taxa known to occur in it. Different taxonomic treatment over time has resulted in the number of endemic species recognised varying from 47 in 1880 to 21 in 1978 and 33 in 2005. The present enumeration of species in the avifaunal list for Sri Lanka is based on the work of the Ceylon Bird Club Rarities and Records Committee, as embodied in Henry (1998) and relative to standard ornithological publications for the region. An authentic list for a country reflects the true diversity of the avifauna within it and contributes to the mapping of correct global distribution especially of widespread or migrating taxa. 46 species of birds in Sri Lanka, including 16 endemic to it, are recognised as Threatened. Key words: Birds, country list, endemic, Threatened, breeding species, migrant species Avifaunal diversity in Sri Lanka Sri Lanka is a large continental land-bridge island, located both in the tropical belt and in the South Asia region. The latter is recognised today as a significant avifaunal entity (Rasmussen & Anderton, 2005). The geographic position and topography of Sri Lanka has given the island three major climatic zones and a wide variety of habitats

within each. These zones and habitats support an avifauna of high species diversity. The majority of these species are breeding residents. Originally from a common landmass shared with India, many have evolved in to distinct forms within the island’s substantial montane massifs which have

TAPROBANICA, ISSN 1800-427X. April, 2009. Vol. 01, No. 01: pp. 28-35. © Taprobanica Nature Conservation Society, 146, Kendalanda, Homagama, Sri Lanka.

28



remained isolated for a long time, despite close proximity and intermittent connection of the island to the mainland (Newton, 2003). The rest are seasonal migrants. As Sri Lanka is at the tip of Peninsular India many species migrating annually from the northern autumn-winter to the tropics along the Central Asian-Indian flyway end their southward journey in this island. Likewise, certain pelagic species which migrate from the southern autumn-winter, northward to the Indian Ocean, occur within the oceanic limits of the country. The current status of avifauna in Sri Lanka is presented on the basis of authentic records of species known from the island. To date these stand at 236 breeding and 203 purely migrant species. By the latter term is meant migrant species with no resident population in the island. The resident species belong to 66 families, and the migrant species to 29 of these and to a further 15 families with no representative resident species (Rasmussen & Anderton, 2005). Most bird species in Sri Lanka are shared with the adjacent Indian mainland and the rest of Asia. The distribution of every resident species in Sri Lanka except those endemic to it extends northward beyond the island either as the same or a different subspecies (Rasmussen, 2005a; Rasmussen & Anderton, 2005). Many migrant species occurring in the Indian mainland also occur in Sri Lanka (Rasmussen & Anderton, 2005). The uniqueness of the avifauna of Sri Lanka is reflected in the diversity of the taxa confined to the island. At present 33 species and 68 subspecies among the other resident species are recognised as endemic to Sri Lanka (Rasmussen & Anderton, 2005). Other than the species endemic to the island, 54 species occurring in it, including six winter migrants, are considered endemic to the South Asia region (Gjershaug et al., 2008; Rasmussen & Anderton, 2005). Changes in records over time After systematic study of the avifauna of Sri Lanka began in the 18th century the diversity of bird species recorded in it increased gradually over the years, with the recognition from time to time of more breeding species and still more migrant species. The major ornithological work for the island in the 19th century featured a total of 371 species (Legge, 1880). This figure has now risen to 439, according to the records shown below.

Most of the increase since then has been due to the identification of new migrant species during the last and present centuries. Four resident species were added to the total during this time. The Blue-eared Kingfisher Alcedo meninting Horsfield, 1821 and Common Coot Fulica atra Linnaeus, 1758 were first recorded in 1894 and 1924 respectively (Whistler, 1944). A species new to science was discovered in 2001, the Serendib Scops Owl Otus thilohoffmanni Warakagoda & Rasmussen, 2004, endemic to Sri Lanka (Clements, 2007; König & Weick, 2008; Warakagoda & Rasmussen, 2004). In 2003 a long-overlooked species was identified from a museum specimen: Marshall’s Iora Aegithina nigrolutea (Marshall, 1876), otherwise known only in India (Rasmussen & Anderton, 2005; Wells et al., 2003). The number of resident species further increased by three as a result of taxonomic revisions among certain closely related taxa, viz. two resident owlets of the genus Glaucidium Boie, 1826 (König et al., 1999; Sibley & Monroe, 1990), two resident Dicrurus Vieillot, 1816 drongos and a group of one resident and two migrant Hirundo Linnaeus, 1758 swallows (Rasmussen & Anderton, 2005). A high endemism in the island’s avifauna has been recognised since the 19th century. By its end 47 species had been recognised as endemic to Sri Lanka amongst a total of 289 resident species as then considered (Legge, 1880). However, in the first half of 20th century the Biological Species Concept was applied in a broad manner to birds in the Indian region (Wijesinghe, 2007). This led to many taxa closely related evolutionarily and isolated geographically which were previously classified as distinct species being treated as subspecies of a smaller number of widespread polytypic species. A result of this practice was a gradual reduction in the number of species endemic to Sri Lanka. This had decreased to 21 by the late 1970s (Phillips, 1978). Since the 1990s, however, ornithologists began gradually re-elevating some populations of distinct taxa thus considered subspecies back to their original species status. This caused a rise again in the number of endemic species in Sri Lanka to 23 (Sibley & Monroe, 1990), 26 (Wijesinghe, 1994) and 33 (Rasmussen & Anderton, 2005). It appears that further revisions in species-level taxonomy may

29



increase the endemic species in Sri Lanka still closer to the number recognised in Legge (1880).

Recording the status of avifauna: a history The birds of Sri Lanka, as mentioned above, first drew attention as a subject for systematic study in the 18th century. This was initiated by the Dutch governor J. G. Loten (Wijesinghe, 1997). Since then the avifauna of the island has attracted great interest, and throughout the British colonial era studies were carried out relating to their diversity, distribution and taxonomy. The results were reflected in Legge (1880), by far the most comprehensive work before the 20th century, and several major publications around the end of that era, viz. Wait (1931), Whistler (1944), Phillips (1953) and Henry (1955).

The last checklist for Sri Lanka with extensive annotation and reflecting the status of these three aspects of its avifauna was Phillips (1978). This was based on the work of two centuries on the birds of the country. It features a total of 392 species. 230 species are recorded as breeding residents, including 21 endemic species and a further 81 endemic subspecies. 162 species are recorded as pure migrants. This was the major reference work on the status of the avifauna of Sri Lanka for two decades until the publication of the third revised edition of G.M. Henry’s popular Guide to the Birds of Sri Lanka (Henry, 1998). The latter work updated species diversity, distribution and taxonomy, with the recognition of 26 species endemic to the island. Its taxonomy was based on Wijesinghe (1994), a checklist with brief annotations, updated to its time. Henry (1998) featured a total of 430 species for the island. Compared with Phillips it recognised one more breeding species, with the ‘splitting’ of two resident taxa of owlets (see above), and 37 new migrant species, according to the decisions of the Ceylon Bird Club Rarities and Records Committee (CBCRRC) (see below).

The current status This paper discusses and presents the status of the species diversity of birds in Sri Lanka based on (a) Henry (1998), and thereafter (b) the subsequent authoritative work of the CBCRRC and (c) the regional works of recognised international ornithologists.

(a) We use Henry (1998) as the basis for our evaluation as it is the most recent national work which records the species accepted into the main country list by the CBCRRC at the time of its publication. (b) The CBCRRC was formed in 1985 (as the CBC Rarities Committee) by the Ceylon Bird Club (CBC), established in 1943 (Ceylon Bird Club Notes, 1985). The club recognised the importance of having such an evaluating committee in line with the practice in other countries where care is exercised to maintain authentic records of bird species occurring within them for their country lists. The chief reason for forming this body was the increase in reports of sightings of new and rare migrant species in Sri Lanka since the 1980s, mainly due to a large increase in birdwatchers and the availability of new literature on bird identification. Identification of such species by sight only (without a specimen) is often found to be erroneous when a high degree of care is not exercised in the field. A less careful observation of what appears to be an unusual bird in the field may well result in a misidentification of species. In order to evaluate reports of sightings of new or rare migrants, submitted to it or published elsewhere, the Committee seeks, inter alia, detailed notes describing important identification features of the species made during observation and preferably a sketch made in the field. Today it is possible to provide further evidence through photographs, video footage and sound recordings if the relevant equipment was available with the observer/s. The other task of the Committee was to look into records, which had already existed in the country list for any length of time, which it regarded as doubtful. The remaining species in the list were those each with at least one specimen held in a museum, authentically known to be from Sri Lanka and of which the identification was not in doubt, and those observed in the field whose identification was beyond doubt in the evaluation of the Committee. Since its inception the Committee has consisted of ornithologists with a high level of knowledge and skill in field observation, and extensive experience in the identification of birds in the field and in museum collections. Several years ago the Committee was broadened to include recognised

30



international experts in the ornithology of the South Asian and Asian regions, to enhance the standard of evaluation. (c) The third source for inclusion of species in the country list is recent publications based on work of regional scope by recognised authorities. Chief among these is Rasmussen & Anderton (2005), the most recent standard work for the region. This was the result of 1½ decades of comprehensive and meticulous work on museum specimens, records of

field observations and audio recordings of vocalisations (Rasmussen, 2005b). The other pertinent works are: Wells et al. (2003), Rasmussen (2005a), Collar (2006) and Gjershaug et al. (2008). Accordingly, subsequent to Henry (1998) the country list has changed by the recognition of six breeding, species, the acceptance or recognition of eight migrant species, and the removal of two species by the CBCRRC. Details are provided in Tables 1 - 3.

Table 01: Breeding species recognised subsequent to Henry (1998)

a These species were already in the country list as migrants. b D. lophorinus was formerly treated as a subspecies of a polytypic resident species D. paradiseus (Linnaeus, 1766). It is treated now as a distinct species. Table 02: Migrant species accepted / recognised subsequent to Henry (1998)

c H. daurica was formerly represented in Sri Lanka by a breeding resident subspecies H. d. hyperythra Blyth, 1849 and two migrant subspecies. The resident taxon is now recognised as a distinct species H. hyperythra, endemic to the island. Thus, H. daurica is now added to the country list as a separate migrant species (in two subspecies). Table 03: Species removed from country list by CBCRRC subsequent to Henry (1998) Nordmann’s Greenshank Tringa guttifer (Nordmann, 1835)d (Ceylon Bird Club Notes, 1998)

Java Sparrow Padda oryzivora (Linnaeus, 1758)e (CBCRRC list: in preparation) d Formerly included as a vagrant species. e Formerly included as an escaped and occasional breeding species.

Indian Spot-billed Duck Anas poecilorhyncha Forster, 1781a (Gunawardena & Sirivardana, 2003)

Bridled Tern Sterna anaethetus Scopoli, 1786a (Perera, 2003)

Serendib Scops Owl Otus thilohoffmanni Warakagoda & Rasmussen, 2004 (Clements, 2007; König & Weick, 2008; Warakagoda & Rasmussen, 2004).

Blue-tailed Bee-eater Merops philippinus Linnaeus, 1766a (Samarasinha, 2003)

Marshall’s Iora Aegithina nigrolutea (Marshall, 1876) (Rasmussen & Anderton, 2005; Wells et al., 2003)

Ceylon Crested Drongo Dicrurus lophorinus Vieillot, 1817b (Rasmussen & Anderton, 2005)

Rufous-necked Stint Calidris ruficollis (Pallas, 1776) (Ceylon Bird Club Notes, 1998)

Pacific Swift Apus pacificus (Latham, 1802) (Ceylon Bird Club Notes, 1998)

Yellow-rumped Flycatcher Ficedula zanthopygia (Hay, 1845) (Ceylon Bird Club Notes, 2004)

Pectoral Sandpiper Calidris melanotos (Vieillot, 1819) (Ceylon Bird Club Notes, 2004)

Bay-backed Shrike Lanius vittatus Valenciennes, 1826 (Ceylon Bird Club Notes, 2004)

Dusky Warbler Phylloscopus fuscatus (Blyth, 1842) (Ceylon Bird Club Notes, 2004)

Common Rosefinch Carpodacus erythrinus (Pallas, 1770) (Ceylon Bird Club Notes, 2004)

Red-rumped Swallow Hirundo daurica Laxmann, 1769c (Rasmussen & Anderton, 2005)

31



This brings the current total of the country list to 439 species, with 236 breeding species, and 203 pure migrants, including rare, very irregular migrants and vagrants. 33 species are now recognised as endemic to Sri Lanka, according to Rasmussen & Anderton (2005). The present authors are aware that reports of several more new migrant species currently await evaluation by the CBCRRC. Of these the ones accepted by this committee will thereby be added to the country list. The authors believe, considering relevant documents perused by themselves, that a further 13 species have been reported with positive evidence, and that it is very probable these will be accepted by the CBCRRC. They believe so with a lesser, but still high, degree of probability, of four more species. On the foregoing the country list may

increase to between 452 and 456 species in the near future.

The importance of correct avifaunal records A list with authentic records reflects the true natural diversity of the avifauna of a country. For widespread or migrating taxa records in such lists form an essential part of the information used to map their global distribution and movement in winter. An incorrect record of the occurrence of such a taxon in Sri Lanka leads to an incorrect idea of its range in these aspects. It is therefore important to maintain criteria of a high standard in the evaluation of reports and previous records when including a migrant species into the country list. Endemic taxa 33 species of birds are recognised as endemic to Sri Lanka (Rasmussen & Anderton, 2005). They are listed in Table 4.

Table 04: Species endemic to Sri Lanka Ceylon Spurfowl Galloperdix bicalcarata (Forster, 1781)

Ceylon Junglefowl Gallus lafayetii Lesson, 1831

Ceylon Woodpigeon Columba torringtonii (Blyth & Kelaart, 1853)

Ceylon Green Pigeon Treron pompadora (Gmelin, 1789)

Ceylon Lorikeet (Hanging-Parrot) Loriculus beryllinus (Forster, 1781)

Layard’s Parakeet Psittacula calthropae (Blyth, 1849)

Green-billed Coucal Centropus chlororhynchos Blyth, 1849

Red-faced Malkoha Phaenicopaeus pyrrhocephalus (Pennant, 1769)

Serendib Scops Owl Otus thilohoffmanni Warakagoda & Rasmussen, 2004

Chestnut-backed Owlet Glaucidium castanonotum (Blyth, 1852)

Ceylon Grey Hornbill Ocyceros gingalensis (Shaw, 1811)

Yellow-fronted Barbet Megalaima flavifrons (Cuvier, 1816)

Ceylon Small barbet Megalaima rubricapillus (Gmelin, 1788)

Crimson-backed Woodpecker (Flameback) Chrysocolaptes stricklandi (Layard, 1854)

Ceylon Swallow Hirundo hyperythra Blyth, 1849

Ceylon Wood Shrike Tephrodornis affinis Blyth, 1847

Black-capped Bulbul Pycnonotus melanicterus (Gmelin, 1789)

Yellow-eared Bulbul Pycnonotus penicillatus Blyth,1852

Spotted-winged Thrush Zoothera spiloptera (Blyth, 1847)

Ceylon Scaly Thrush Zoothera imbricata Layard, 1854

Ceylon Whistling-Thrush Myophonus blighi (Holdsworth, 1872) Dusky-blue Flycatcher Eumyias sordidus (Walden, 1870)

32



Ashy-headed Laughing-Thrush Garrulax cinereifrons Blyth, 1852

Brown-capped Babblerf Pellorneum fuscocapillus (Blyth, 1849)

Ceylon Scimitar Babblerf Pomatorhinus melanurus Blyth, 1847

Ceylon Rufous Babbler Turdoides rufescens (Blyth, 1847)

Ceylon Bush Warbler Elaphrornis palliseri (Blyth, 1852)

Legge’s Flowerpecker Dicaeum vincens (Sclater, 1872)

Ceylon White-eye Zosterops ceylonensis Holdswoth, 1872

White-headed(-faced) Starling Sturnia albofrontata (Layard, 1854)

Ceylon Hill-Mynah Gracula ptilogenys Blyth, 1846

Ceylon Crested Drongo Dicrurus lophorinus Vieillot, 1817

Ceylon Blue Magpie Urocissa ornata (Wagler, 1829) f These two species each comprises two subspecies. The other species are monotypic.

68 subspecies of birds are recognised as endemic to Sri Lanka among the species resident in the island other than the endemic species (Rasmussen & Anderton, 2005). 55 of them are each differentiated from one or more closely related subspecies in the Indian mainland by biometrics and/or mere shade of plumage colour. The other 13 are more distinct than this from such closely related taxa by differing in biometrics and in colour/s of discrete elements in

plumage. In the opinion of Wijesinghe (1994), Rasmussen & Anderton (2005) and the present authors five taxa in this last group are adequately distinct from such related subspecies in biometrics and/or coloration of plumage they may reasonably be considered for taxonomic revision elevating them to species status. These are listed in Table 5.

Table 05: Endemic taxa which may be elevated to species status Red-backed Wooodpecker (Black-rumped Flameback) Dinopium benghalense psarodes (Lichtenstein, 1793)

Ceylon (Asian) Paradise Flycatcher Terpsiphone paradisi ceylonensis (Zarudny & Härms, 1912)

Indian Blackbird Turdus simillimus kinnisii (Kelaart, 1851)

Black-(Dark-)fronted Babbler Rhopocichla atriceps nigrifrons (Blyth, 1849) / R. a. siccata Whistler, 1941

Ceylon Hill (Black-throated) Munia Lonchura kelaarti kelaarti (Jerdon, 1863) Threatened species According to The 2007 Red List of Threatened Fauna and Flora of Sri Lanka (IUCN & MENR, 2007) 46 bird species in Sri Lanka are threatened, and these include 16 endemic species. Numerical summary Table 06: Status of avifauna in Sri Lanka as at December 2008

Note to table: See also the last paragraph in the section ‘The current status’. Acknowledgements We thank the Editor of this journal, Thasun Amarasinghe, for the invitation to write this paper, the Committee of the Ceylon Bird Club for

Subspecies endemic to country of other resident species 68

Threatened species:

Breeding resident 46

Endemic (among the above) 16

Breeding species 236

Purely migrant species 203

Total number of species in country list 439

Species endemic to country 33

33



permission to use unpublished information in the possession of the club, Priyantha Wijesinghe for providing certain taxonomic authorships and Thilo Hoffmann, Pathmanath Samaraweera and Kithsiri Gunawardena for their valuable comments on a draft of this paper. Literature Cited Ceylon Bird Club Notes, 1985. Rarities Committee: 26. Ceylon Bird Club Notes, 1998. Records accepted by CBC Rarities Committee: 53-54. Ceylon Bird Club Notes, 2004. Ceylon Bird Club Records Committee decisions, 109-110. Clements, J. F., 2007. The Clements Checklist of the Birds of the World. 6th Edition. Christopher Helm, London: 864. Collar, N. J., 2006. A partial revision of the Asian babblers (Timaliidae). Forktail, 22: 85-112. de Silva Wijeyeratne, G., 2008. Birds of Sri Lanka: A Pictorial Guide and Checklist. Jetwing Eco Holidays, Colombo: 66. de Silva Wijeyeratne, G. and D. Warakagoda, 2001. A Checklist of the Birds of Sri Lanka. Sri Lanka Natural History Society and Jetwing Research Initiative, Colombo: 24. Gjershaug, J. O., O. H. Diserud, P. C. Rasmussen and D. Warakagoda, 2008. An overlooked threatened species of eagle: Legge’s Hawk Eagle Nisaetus kelaarti (Aves: Accipitriformes). Zootaxa, 1792: 54-66. Gunawardena, K. and U. Sirivardana, 2003. Report in: Ceylon Bird Club Notes: 39 Harrison, J. and T. Worfolk, 1999. A Field Guide to the Birds of Sri Lanka. Oxford University Press, Oxford: 219. Henry, G. M., 1955. A Guide to the Birds of Ceylon. Oxford University Press, London: 445. Henry, G. M., 1998. A Guide to the Birds of Sri Lanka. Third Edition: revised & enlarged by T. W. Hoffmann, D. Warakagoda and U. Ekanayake. Oxford University Press, London and K. V. G. de Silva & Sons, Kandy: 488. IUCNSL and MENR, 2007. The 2007 Red List of threatened Fauna and Flora of Sri Lanka. Colombo, IUCN Sri Lanka: 148.

König, C. and F. Weick, 2008. Owls of the World. Second Edition. Christopher Helm, London: 528. König, C., F. Weick and J. -H. Becking, 1999. Owls: A Guide to the Owls of the World. Pica Press, Sussex: 462. Kotagama, S. W., R. I. De Silva, A. S. Wijayasinha and V. Abeygunawardane, 2006. Avifaunal List of Sri Lanka. In: Bambaradeniya, C. N. B. (Ed.). The Fauna of Sri Lanka: Status of Taxonomy, Research and Conservation. The World Conservation Union and Government of Sri Lanka, Colombo: 164-203. Legge, W. V., 1880. A History of the Birds of Ceylon. (1983 Edition) Vols. 1-4. Tisara Prakasakayo Ltd. Dehiwala: 1882. Newton, I., 2003. The Speciation and Biogeography of Birds. Academic Press. London: 668. Perera, L., 2003. Report in: Ceylon Bird Club Notes: 127 Phillips, W. W. A., 1953. A (1952) Revised Checklist of Birds of Ceylon. The National Museums of Ceylon, Colombo: 132. Phillips, W. W. A., 1978. Annotated Checklist of the Birds of Ceylon (Sri Lanka). Revised Edition. Wildlife and Nature Protection Society of Ceylon in association with The Ceylon Bird Club, Colombo: 92. Rasmussen, P. C., 2005a. Biogeographic and conservation implications of revised species limits and distributions of South Asian birds. Zoologische Mededelingen Leiden, 79-3 (13): 137-146. Rasmussen, P. C., 2005b. On producing Birds of South Asia. Indian Birds, 1(3): 50-56. Rasmussen, P. C. and J. C. Anderton, 2005. Birds of South Asia: The Ripley Guide. Vols. 1 and 2. Smithsonian Institution and Lynx Edicions, Washington, D.C. and Barcelona: 378, 683. Ripley, S. D., 1982. A Synopsis of the Birds of India and Pakistan: together with those of Nepal, Bhutan, Bangladesh and Sri Lanka. Bombay Natural History Society, Bombay: 652. Samarasinha, R., 2003. Report in: Ceylon Bird Club Notes: 91-92 Sibley, C. G. and B. L. Monroe, 1990. Distribution and Taxonomy of Birds of the World. Yale University Press, New Haven: 1111.

34



Wait, W. E., 1931. Manual of the Birds of Ceylon. Second Edition. The Director, Colombo Museum, Colombo, 494. Warakagoda, D. H. and P. C Rasmussen, 2004. A new species of scops-owl from Sri Lanka. Bulletin of the British Ornithologists’ Club, 124 (2): 85-105. Wells, D. R., E. C. Dickinson and R. W. R. J. Dekker, 2003. Sytematic notes on Asian birds. 34. A preliminary review of the Aegithinidae. Zoologische Verhandelingen Leiden, 344: 7-15. Whistler, H., 1944. The Avifaunal Survey of Ceylon, conducted jointly by the British and Colombo Museums. Spolia Zeylanica, 23 (3 & 4): 119-322. Wijesinghe, D. P., 1994. Checklist of the Birds of Sri Lanka. Ceylon Bird Club, Colombo, 49. Wijesinghe, D. P., 1997. Bird Study in Sri Lanka: an historical perspective. Bulletin of Oriental Bird Club, 26: 26-31. Wijesinghe, D. P., 2007. Comments on Birds of South Asia: The Ripley Guide by Pamela C. Rasmussen and John C. Anderton (2005). Ceylon Bird Club Notes: 83-95. Appendix: An overview of publications subsequent to Henry (1998) recording the avifauna of Sri Lanka After Henry (1998) there have been four publications in English which list the birds of the country. As they are few they are reviewed here. 1) A Field Guide to the Birds of Sri Lanka (1999) by J. Harrison and T. Worfolk. This is an excellent field guide, which has proved its value in bird identification and is widely used. It features all the species accepted by the CBCRRC to the above time. Its major flaw is that it includes (illustrates and describes) species whose occurrence in the island still needs confirmation (even to the time of this paper), and which were therefore listed outside the authentic country list by the CBCRRC, as reflected in Appendix I of Wijesinghe (1994). That gives an incorrect impression to most users that there are authentic records of the occurrence of these birds in Sri Lanka. This has led some observers to misidentify as these species individuals of other species with a similar appearance, which occur regularly in Sri Lanka.

2) A Checklist of the Birds of Sri Lanka (2001) by G. de Silva Wijeyeratne and D. Warakagoda. This is designed in the form of a ‘tick list’ of species and subspecies of the birds of Sri Lanka to mark those noted in the field by a reader. The list comprises the species recorded authentically to that date as published in Henry (1998), those additionally accepted by the CBCRRC and six others, which in the opinion of its authors have been recorded reliably. A total of 434 species is listed. The six species mentioned above were all subsequently accepted into the country list by the CBCRRC. 3) Avifaunal List of Sri Lanka by S. W. Kotagama et al. in The Fauna of Sri Lanka: Status of Taxonomy, Research and Conservation (2006). This is an enormously inflated list of species, which gives a corresponding idea of the diversity of the avifauna of Sri Lanka, with 482 species included. It lists many species from reports not considered authentic in many ornithological works dating back to the early 20th century and thus not included in them. Several other species are listed on the basis of reports which had not been accepted by the CBCRRC, and reports whose contents fall well short of the level of authenticity required by a standard rarities committee. A clearly inadequate criterion is used for accepting sight records of rare migrants: viz. merely being “confirmed by more than one observer” as an alternative to a proper published description. Such defects have even drawn into this list, from a report in the last decade, a species known to date to be endemic to Indochina and its vicinity. 4) Birds of Sri Lanka: A Pictorial Guide and Checklist (2008) by G. de Silva Wijeyeratne This is similar to de Silva Wijeyeratne & Warakagoda (2001) in its format and the basis used to list species, with the addition of photographs of a large number of these. It deals with reports to May 2007. The total number of species presented is 444.

35

BOTEJUE, 2009


AN OBSERVATION OF Vanellus indicus BODDAERT, 1783 (AVES: CHARADRIIDAE) FEED ON AN EXOTIC Laevicaulis altae FERUSSAE, 1821 (GASTROPODA: VERONICELLIDAE) AT A HUMAN HABITATION IN SRI LANKA


W. Madhava S. Botejue

Taprobenica Nature Conservation Society, 146, Kendalanda, Homagama, Sri Lanka E-mail: [email protected] Abstract Red–Wattled Lapwing is a common resident bird distributed throughout Sri Lanka. Laevicaulis altae is an exotic land slug found in home gardens and cultivations in the lowland wet zone and intermediate zone. L. altae and other exotic snails and slugs are pests, which is a serious and growing problem in Sri Lanka. This observation describes the behaviour of Red–Wattled Lapwing feeding on the exotic land slug at the premises of the Open University of Sri Lanka, Colombo. Key Words: Red–Wattled Lapwing, Laevicaulis altae, exotic slug, feeding behaviour, predation, Sri Lanka. Introduction Red–Wattled Lapwing, Vanellus indicus Boddaert, 1783 is a common resident bird distributed throughout the edges of tanks, dry paddy fields and open lands near water bodies in the low country and lower hills of Sri Lanka (Ali & Ripley, 1980; Harrison, 1999; Henry, 1998; Legge, 1983). This bird is about the size of a domestic pigeon with a black, white and bronze-brown coloration and long, slender yellow legs. The most distinguishable character of this bird is the fleshy red wattle in front

of the eye and the red ring around the eye. Sexes are alike (Ali & Ripley, 1980; Harrison, 1999; Henry, 1998; Legge, 1983). This species take most of its food on dry ground, which consists of ground dwelling insects like beetles, grasshoppers, termites and ants, insect larvae like caterpillars and worms, mollusks, aquatic insects as well as vegetable matter (Ali & Ripley, 1980; Henry, 1998; Legge, 1983). It feed casually during the day, but feeds more actively in the early mornings, evenings and


36

AN OBSERVATION OF Vanellus indicus FEED ON AN EXOTIC Laevicaulis altae AT A HUMAN HABITATION OF SRI LANKA


throughout the night, especially when moonlight (Ali & Ripley, 1980; Harrison, 1999). Laevicaulis altae Ferussae, 1821 is an exotic land slug found in home gardens and cultivations in the lowland wet zone and in the wetter parts of intermediate zone (Raheem & Naggs, 2006; Priyadarshana, 1998). This species prefers shady damp places, often seen around wells and ponds, on slippery stones and under damp dead leaves (Priyadarshana, 1998). On 14th May 2008 between 11:35 hr and 11:55 hr I observed a mature Red – Wattled Lapwing feeding on an exotic land slug L. altae at the premises of the Open University of Sri Lanka (OUSL). OUSL (6o 53’ 10” N and 79o 52’ 60” E; alt. 10 m) is situated about 6 km from the Colombo city in Colombo District in Western Province of Sri Lanka. The vegetation resembles home gardens and the “Kirulapona” canal runs along the west, northwest border of the location. The ground was covered with grass and the canopy cover was about 20%. The observations were made by the naked eye without any disturbances to the bird. First the bird was observed on the ground foraging about 5 m away from me. The temperature and the humidity at that time were 31 oC and 76 % respectively. The weather was sunny. At once the bird dipped forward steeply and caught some thing, which was on the grass. Then it moved about 3 m towards me in an irregular zigzag manner. It was holding a large L. altae crosswise at the tip of the bill (Fig. 01). Then the bird kept the slug on the ground and looked around vigilantly for about 20 seconds. After that the bird rubbed the slug on the grass using its bill for about 1 minute, looking up at regular intervals. Then it again took up the slug in the bill and looked around vigilantly for about another 2 minutes. After that it moved away about 2 m and kept the slug on the ground. Then the bird rubbed the slug on the grass for about another 2 minute, looking around at regular intervals. Likewise it repeated the same action for three times. Then the bird moved about another 1 m and swallowed the slug. Then the bird foraged the surrounding about 5 minutes until a domestic dog disturbed the bird. Hence the bird ran about 5m and flew away. According to my observation Red–wattled Lapwings are seemed to be unaccustomed in feeding on slugs. The bird seemed to have some hesitation about feeding on slugs, because it took

about 15 minutes to devour a L. altae. I assume that the reason for the bird to rub the slug on the grass is to wipe off the mucous produce by it. Because the mucous produced by slugs may have a defensive function. It may be distasteful to the predator or it may adhere to the mouthparts of the predator and may thus hamper its attack on the slug (Dinarzarde Raheem, August 2008, personal communication). According to Richter, 1980 in Ariolimax columbianus the caudal mucus plug covers the caudal pit which serves as a defense mechanism. Therefore I assume that L. altae also demonstrate the same mechanism. The previous literature also explains the vigilant behaviour of the Red – wattled Lapwing (Ali & Ripley, 1978; Henry, 1998). L. altae and other exotic snails and slugs are pests for native biodiversity, which is a serious and growing problem in Sri Lanka (Naggs & Raheem, 2002; Raheem & Naggs, 2006). If the Red – wattled Lapwing is a common predator of the L. altae, it will be helped to control the spreading of this introduced slug. Fig. 01: Red–wattled Lapwing is holding a large L. altae crosswise at the tip of the bill.

Acknowledgements I wish to thank Sujan Henkanaththegedara, M. M. Bahir and anonymous reviewers for their valuable comments and reviewing the manuscript. I would like to thank Dinarzarde Raheem (BMNH) for valuable comments and helped improve the document. I also wish to thank my colleagues at the OUSL for supporting the field observation.

Literature Cited Ali, S. and S. D. Ripley, 1980. Handbook of the Birds of India and Pakistan, Vol. 02, Second edition. Oxford University Press, New Delhi: 347.

37

BOTEJUE, 2009


Harrison, J. and T. Worfolk, 1999. A Field Guide to the birds of Sri Lanka. Oxford University Press, Oxford: 219. Henry, G. M., 1998. A guide to the birds of Sri Lanka, Third Edition: revised & enlarged by T. W. Hoffmann, D. Warakagoda and U. Ekanayake. Oxford University Press, London and K. V. G. de Silva & Sons, Kandy: 488. Legge, W. V., 1883. A history of the birds of Ceylon, Second edition. Tisara Prakashakayo, Dehiwala, Sri Lanka. 1624. Naggs, F. and D. Raheem, 2002. Sri Lankan snails. The Natural History Museum, London, UK. Priyadarshana, T. G. M., 1998. Land Slugs of Sri Lanka (Phylum Mollusca, Family Veronicellidae). Sri Lanka Naturalist, 2 (1&2): 14-15. Raheem, D. and F. Naggs, 2006. An Illustrated Guide to the Land Snails of Sri Lankan Natural Forest and Cultivated Habitats. The Natural History Museum, London: 6. Richter, K. O., 1980. Movement, defense and nutrition as functions of the caudal mucous plug in Ariolimax columbianus. Veliger, 23: 43-47.

38

EDIRISINGHE AND AMARASINGHE, 2009


AN UNUSUAL MISLEAD COMMUNICATION BEHAVIOUR OF

Duttaphrynus melanostictus (SCHNEIDER, 1799) (AMPHIBIA: BUFONIDAE)

AND Polypedates cruciger BLYTH, 1852 (AMPHIBIA: RHACOPHORIDAE)

AT A HUMAN HABITATION IN SRI LANKA


W. Gayan M. Edirisinghe 1 and A. A. Thasun Amarasinghe

2,3

1 The Young Zoologists' Association of Sri Lanka, Department of National Zoological Gardens, Dehiwala, Sri

Lanka. 2 Taprobanica Nature Conservation Society, 146, Kendalanda, Homagama, Sri Lanka.


Abstract Duttaphrynus melanostictus and Polypedatus cruciger are common amphibians in Sri Lanka, being

sympatric in many habitats, while P. cruciger is endemic to the island. This is the first published observation

on an unusual mislead communication behavior on D. melanostictus and P. cruciger at a human habitat in Sri Lanka. Considering the field observation, we can assume both D. melanostictus and P. cruciger males

may accidentally catch almost immediately without identifying each other properly in their breeding periods.

We suggest that these amphibians communicate with their unique call but their eyesight is not developed to

identify its partner easily.

Key Words: Duttaphrynus melanostictus, Polypedatus cruciger, unusual communication behavior, Sri Lanka

Introduction There are 109 species of amphibians in Sri Lanka

and 92 of them are endemic. Also further new

species will be described in the future

(Meegaskumbura et al., 2002). These 109 species

are classified under seven families; Ichthyophiidae,

Bufonidae, Dicroglossidae, Microhylidae, Ranidae,

Nyctibatrachidae and Rhacophoridae (Frost et al.,

2006). Family Bufonidae is one of the most

common amphibian families in south and Southeast

Asia (Pough et al., 2004). Bufonidae consists of



39

AN UNUSUAL MISLEAD COMMUNICATION BEHAVIOUR OF Duttaphrynus melanostictus AND Polypedates cruciger


three genera Bufo, Duttaphrynus and Adenomus, the

genus Duttaphrynus consists of two species and

among them one species is endemic to Sri Lanka

(Frost, et al., 2006; Manamendra-Arachchi &

Pethiyagoda, 1998; Manamendra-Arachchi & Pethiyagoda, 2006).

Duttaphrynus melanostictus is distinguished from

all the other Sri Lankan Duttaphrynus species by

having parietal ridges absent and two longitudinal

rows of large warts which distributed all length of

inter parotid area. Snout-vent length of mature

males is 50.3-90.0 mm, gravid females 70.0-95.0

mm (Manamendra-Arachchi & Pethiyagoda, 1998;

Manamendra-Arachchi & Pethiyagoda, 2006). The

Family Rhacophoridae consists of 69 species

belonging to three genera: Philautus, Polypedates

and Rhacophorus. There are 5 species of genus

Polypedatus in Sri Lanka. Four of them are endemic

to the country include P. cruciger (Dutta &

Manamendra-Arachchi, 1996; Frost, et al., 2006).

Polypedatus cruciger, which is endemic to Sri

Lanka, is distinguished from all the other Sri

Lankan Polypedatus species by having the calcar absent and possessing a co-ossified skull. Snout-

vent length of mature males is 50.0-59.8 mm,

gravid females 72.0-90.0 mm (Dutta &

Manamendra-Arachchi, 1996; Manamendra-

Arachchi & Pethiyagoda, 2006). This short

communication is intended to provide information

on an observation relating to unusual mislead communication behavior on Duttaphrynus

melanostictus and P. cruciger at a human habitat in

Sri Lanka. Both species are common in Sri Lanka

and also sympatric in many habitats. This is the first

record of an unusual mislead communication

behavior involving D. melanostictus and P.

cruciger.

Observation The observations were made in Kesbewa (alt. 10 m

a.s.l.) in Colombo district in Western province of

Sri Lanka. The geographical coordinates of the

study area are 06o 40’ 54” N and 80o 00’ 34” E,

approximately 200 m away from Kesbewa town.

The observations were made by the naked eye, 2 m

away from the toad and the tree-frog from 09:15 hr. to 10:15 hr. No disturbances were made to animals

during the time of observation. A mature male

Duttaphrynus melanostictus was observed on 29th

March 2006 at about 09:15 hr., while it was puffing

its vocal sac and calling sharp on the ground. The

temperature and the humidity at that time was 29 oC

and 79%. The weather was shady. Firstly the male

D. melanostictus jumped towards an artificial pond

up to about one meter to the edge of the pond. At

once a mature male Polypedatus cruciger jumped

on to toad’s back (Fig. 01).

Fig. 01: The tree-frog jumped to toad’s back

The male tree-frog stayed on the male toad’s back

for about ten minutes. During this time the toad

tried to escape from the tree-frog. Finally the toad

kicked the tree-frog and then the tree-frog jumped to in front of the toad on the ground (Fig. 02).

Following, the toad jumped forward and caught the

tree-frog tightly. Likewise, the toad stayed on the

tree-frog’s back for 5 minutes (Fig. 03). Afterwards,

the tree-frog escaped from the toad and jumped

again to its back. At this time the tree-frog stayed

on the toad’s back for about another five minutes. During this time, a female P. cruciger jumped near

the unmatched couple (Fig. 04). Immediately the

male toad jumped over and caught the second

female tree-frog (Fig. 05) and gripped it tightly for

about five minutes. After that, the toad released its

grip and escaped. Then the tree-frog jumped into

the pond and disappeared.

Fig. 02: The tree-frog jumped to in front of the toad

40

EDIRISINGHE AND AMARASINGHE, 2009


Fig. 03: The toad jumped forward and caught the tree-

frog tightly

Fig. 04: the tree-frog escaped from the Toad, jumped

again to its back and another tree-frog arrived

Fig. 05: The toad jumped over and caught the second

female tree-frog

Discussion According to Emerson (2001), Giacoma and

Castellano (2001) and Pough et al. (2004) there are

no records of that unusual type of mislead communication between two amphibian species

belonging to two different families. This is the first

documented observation of an unusual mislead

communication behavior involving Duttaphrynus

melanostictus and Polypedates cruciger in Sri

Lanka. We assume this was a mistaken

communication, which was made by both species.

This observation demonstrates that this two species

are clearly attracting their partners by calling,

though both callings were made in the same time

period. Both males of these species expected a female of their species but only a female P. cruciger

came. So when the male P. cruciger spotted the D.

melanostictus it perhaps recognized visually it as its

mate. Therefore the male P. cruciger jumped to

toad’s back.

We could see the toad was not like this because it is

a male. After dispatch Polypedates cruciger male

from its back the male Duttaphrynus melanostictus

saw the female P. cruciger and made the mistake

again and grabbed it as its mate. After few minutes

it may have realized the mistake and gave the

female P. cruciger a chance to escape. We suggest

these tree-frogs communicate with their unique call

but their eyesight is not developed to identify its

partner easily. After considering all the observations

we could conclude that both D. melanostictus and

P. cruciger males may catch each other

immediately without identifying themselves

properly in their breeding periods.

Acknowledgements The authors wish to thank Enrique La Marca, Tzi Ming Leong and Mohomed M. Bahir for useful

comments. And also we would also like to thank

Suranjan Karunarathna (IUCN) and Dinesh

Gabadage (TNCS) for their supports.

Literature Cited Dutta, S. K. and K. N. Manamendra-Arachchi, 1996. The Amphibian Fauna of Sri Lanka. Wildlife Heritage

Trust of Sri Lanka: 230.

Emerson, S. B., 2001. Male advertisement calls:

behavioral variation and physiological processes. In: Ryan, M.J. (Ed.). Anuran Communication.

Smithsonian institution press, Washington and

London: 36-44.

Frost, D. R., T. Grant, J. Faivovich, R. H. Bain, A. Haas, C. F. B. Haddad, R. O. de Sá, A. Channing, M.

Wilkinson, S. C. Donnellan, C. J. Raxworthy, J. A.

Campbell, B. L. Blotto, P. Moler, R. C. Drewes, R. A.

Nussbaum, J. D. Lynch, D. M. Green and W. C.

Wheeler, 2006. The Amphibian Tree of Life. Bulletin of the American Museum of Natural History, 297: 1-

370.

Giacoma, C. and S. Castellano, 2001. Advertisement

call variation and speciation in the Bufo viridis

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AN UNUSUAL MISLEAD COMMUNICATION BEHAVIOUR OF Duttaphrynus melanostictus AND Polypedates cruciger


complex. In: Ryan, M. J. (Ed.). Anuran

Communication. Smithsonian institution press, Washington and London: 205-219.

Manamendra-Arachchi, K. and R. Pethiyagoda, 1998.

A synopsis of the Sri Lankan Bufonidae (Amphibia:

Anura) with description of two new species. Journal of South Asian Natural History, 3 (2): 213–248.

Manamendra_Arachchi, K. and R. Pethiyagoda, 2006. Sri Lankawe Ubhayajeevin “Amphibians of Sri

Lanka” (text in sinhala). Wildlife Heritage Trust of Sri Lanka: 440+88.

Meegaskumbura, M., F. Bossuyt, R. Pethiyagoda, K. Manamendra-Arachchi, M.M. Bahir, M.C.

Milinkovitch and C.J. Schneider, 2002. Sri Lanka: an amphibian hotspot. Science: 298: 379.

Pough, F. H., R. M. Andrews, J. E. Cadle, M. L. Crump, A. H. Savitzky and K. D. Wells, 2004.

Herpetology, Third Edition. Pearson Prentice Hall,

USA: 726

42

SOMAWEERA AND SOMAWEERA, 2009


AN OVERVIEW OF SRI LANKAN SEA SNAKES WITH AN ANNOTATED CHECKLIST AND A FIELD KEY

Submitted: 12 November 2008, Accepted: 15 January 2009

Ruchira Somaweera 1 and Nilusha Somaweera 2

1 Reptile Ecology Lab, School of Biological Sciences, The University of Sydney, Australia. 2 University of Sydney Tropical Ecology Research Facility, Northern Territory, Australia. 1 Corresponding author: [email protected] Abstract Sea snakes of Sri Lanka has not received much attention and the last few decades did not see any unprecedented increase in interest in these largely ignored (in Sri Lanka) snakes, despite heavy attention has been made on certain other groups of herpetofauna. Hence, a new checklist and an overview in ‘general literature’ are much needed. This contribution provides an enumeration of the recorded species, together with an overview of the natural history of the group and a set of field keys for species found in Sri Lankan waters. Keywords: sea snakes, sea kraits, natural history, distribution, identification key, Sri Lanka Introduction Sea snakes: Snakes of at least five distinct lineages inhabit the marine environment. These comprise the file snakes (family Acrochordidae), the mud snakes (family Colubridae: subfamily Homalopsinae), the water snakes (family Colubridae: subfamily Natricinae), the sea kraits (family Laticaudidae) and the true sea snakes (family Hydrophiidae) (Heatwole, 1999), though the term ‘sea snake’ mainly refers to the sea kraits and the true sea snakes. Wall (1909) in the first monograph on sea

snakes identified 11 genera and followed Boulenger (1986)’s classification where all sea snakes were placed under subfamily Hydrophiinae in family Elapidae. Subsequently Smith (1926) in his monograph on sea snakes grouped all sea snakes under family Hydrophiidae which he further divided into two subfamilies: Hydrophiinae and Laticaudinae with Laticauda as the most primitive genus in the latter. All subsequent authors including Dowling (1959 & 1967), Underwood (1967 &


43



1979) and McDowell (1967 & 1974 etc.) followed Simth’s classification with two subfamilies till Burger & Natsuno (1974) erected two separate families for the sea snakes, Laticaudidae (sea kraits) and Hydrophiidae (sea snakes) which was followed later by many other workers till to-date. However, some experts of today consider that the Laticaudids and Hydrophids evolved from different terrestrial representatives of the family Elapidae. Moreover, Rasmussen (1997) showed that the Hydrophids can be separated into two quite different groups, indicating that sea snakes may have evolved three times from terrestrial elapids and provided a phylogeny for the sea snakes. However, this paper follows the ‘two separate families’ classification and the word ‘sea snake’ has been used commonly to denote both Laticaudids and Hydrophids.

History of investigations in Sri Lanka: The first information about the species composition of sea snakes of Sri Lanka was published by Wall (1921) who considered the sea snake wealth to be 22 species belonging to 16 genera but he included all the species in the Indian Ocean around Sri Lanka, not necessarily the species found within Sri Lankan waters. Deraniyagala (1949) identified 19 species but subsequently revised the list to 12 species (Deraniyagala, 1955). Both de Silva (1980) and de Silva (1990) recognised 13 species. Due to the vast limit of the territorial waters of the island, de Silva (1994) suspected the occurrence of Hydrophis fasciatus fasciatus, Hydrophis caerulescens, Hydrophis nigrocinctus and earlier Lapemis hardwicki (now L. curtus) in Sri Lankan waters and listed them. Some of these species have been also mentioned in earlier works and Sri Lanka lies within the reported range of these species. Additionally they have been recorded from the adjacent Madras coasts (Murthy, 1977). However, since there were no confirmed records of these species in Sri Lankan waters, Das & de Silva (2005), de Silva (2006) and Somaweera (2006) did not listed them in the checklists. Das & de Silva (2005) included Laticauda colubrine in the checklist to increase the total up to 14 species. Though Laticaudids have not been recorded from a coast in Sri Lanka, Wall (1921) also included Sri Lanka in its distribution. Somaweera (2006) further included Hydrophis mamillaris in the list, a species most previous author considered to be a synonym of Hydrophis fasciatus but probably represent a distinct species (Wall, 1921; Arne Redsted Rasmussen, pers. comm.; A. Lobo, pers. comm.).

General Remarks on Natural History: The sea kraits of family Laticaudidae inhabit the tropical waters of the western Pacific and northern Indian Oceans (Cogger, 2000) and currently eight species are recognised in the family (Cogger & Heatwole, 2006). Kharin & Czeblukov (2006) splitted the species into two genera, Laticauda and Pseudolaticauda but both Heatwole et al. (2005) and Cogger & Heatwole (2006), refused Kharin’s and Czeblukov’s proposel to make two genera in the family and both papers added all species to the genus Laticauda. The taxonomy of sea snakes belonging to family Hydrophiidae is still not fully resolved, but a total of 58 distinct species are cumulatively identified in literature (Uetz & Hallermann,1995-2008). However, researchers have not reached consensus on the integrity of some taxa (Guinea, 2003). Sea snakes occur in the tropical and subtropical waters of the Indian and Pacific oceans from the east coast of Africa to the Gulf of Panama (Rasmussen, 2000), but were considered to be absent from the Atlantic Ocean (Cozzi, 1980) till Branch (1998) extended the distribution of Pelamis platurus into Atlantic. Sea snakes mostly inhabit shallow waters in the continental shelf but species like Pelamis platurus are pelagic. Contrastingly certain species are know to ascend into rivers and three species, Hydrophis semperi in Phillipines, Hydrophis sibauensis in Indonesia and Laticauda crockeri in Solomon’s islands are found in freshwater rivers and lakes (Cogger et al., 1987; Herre, 1942; Rasmussen et al., 2001). Hydrophis sibauensis which is living more than 100 miles from the sea in a small river system has been never recorded in salt or brackish waters. Furthermore Ineich (1996) recorded the occurrence of “sea snakes” in the Great lake of Cambodia. Sea snakes exhibits dramatic specialisations for their “fully” aquatic habits: a vertically flattened paddle-like tail for propulsion (absent in all other snakes, including other freshwater and brackishwater species); dorsally positioned nostrils each with a valve, salt regulating glands (including lacrimal glands that remove salt as tears), and a single lung that extends nearly the full length of the body (Dunson, 1975). Aipysurus laevis is reported to has a tail with cutaneous photoreceptive ability (light-sensitive) tail which is used to detect approach of predators (Zimmerman & Heatwole, 1990). The Sri Lankan species range in size from ~75 cm Pelamis platurus to the ~300cm Hydrophis spiralis, which is probably the longest sea snake in the world (Fichter, 1982). However Astrotia stokesii with a midbody girth of up to 26 cm at a length of 1.8 m is probably the largest or stoutest of all sea

44



snake (Green, 1997). All sea snakes are air-breathers and a single breath may last up to two hours in the case of Pelamis platurus but usually lasts as little as 30 minutes when the snakes are actively foraging (Heatwole, 1999). All Sri Lankan sea snakes are piscivorous predators that hunt during the day; at dawn or at night and few species eat fish eggs. The yellow-bellied sea snake, Pelamis platurus, is considered to persist the ability to catch fish in open water (Kropach, 1975). Guinea (2003) reported that all the other species corner their prey in crevices or burrows but he does not explain how species like Enhydrina schistosa which prey on Catfish (Voris et al., 1978) and Hydrophis ornatus which is a generalist feeding on many different fish species not only living on the bottom (Rasmussen 1989) catch their prey. Behavioural studies indicate that Pelamis platurus are indeed even sensitive to water motions caused by swimming fish (Heatwole, 1999). To find their mobile prey in often turbid waters olfactory and visual cues may not be the best and only option and sea snakes may also use cutaneous mechanoreceptors and/or inner ear receptors to detect weak water motions such as those generated by prey objects (Westhoff et al., 2005). The genus Laticauda is oviparous thus comes ashore to lay eggs while all other sea snakes are viviparous and give birth to live young in the ocean. A considerable number of species move upstream in rivers to give birth (as mentioned by Porter et al., 1997 etc.). Species like Astrotia stokesii are generally known to have relatively large brood sizes (Heatwole, 1997) and Enhydrina schistosa has large broods in Australian waters (Fry et al., 2001) but there’s only about 4-11 young in an average clutch in Asian waters (Voris & Jayne, 1979). Sea snakes are considered to be breeding annually and the reproductive seasonality varies amongst the sea snakes. A study by Rasmussen (1989) indicates the occurrence of synchronised annual reproductive cycles in Hydrophis ornatus. The gestation periods are lengthy and may vary between six and seven months (Heatwole, 1997). According to Voris & Jayne (1979) mortality amongst the young is high in sea snakes with an estimated 10 to 20% of young Enhydrina schistosa surviving the first year and only 6% of females of this species surviving to reproduce. Distribution: The coastal and deep sea waters of Sri Lanka harbours lot of ideal habitats for sea snakes, including coral reefs, estuaries, bays, lagoons, mangrove forests, sea grass beds, large rivers, canal mouths etc. Fernando & Goonaratne (1983) stated

that sea snakes are common in Palk Bay and off Mullativu. However a vast majority of the sea snake records are from those entangle in nets of fishermen. Hence, most of the locality data coincide with the fishing beds and locations of fishing harbours. The region of marine resources exploitation is mainly confined to the narrow continental shelf, which rarely exceeds 40 km and averages around 22 km in width. The total area of the shelf is about 30000 km2, which is around 6% of the total area of the Exclusive Economic Zone (EEZ) of Sri Lanka (Haputhantri et al., 2008). Hence the recorded species are principally from the continental shelf and deep sea pelagic species have been largely overlooked. Checklist: By and large, we are following the list and systematic given by Somaweera (2006) from family level down. The systematic in this paper are simplified with only family, genus and species levels being considered. They are presented in purely alphabetical order of the family, genus and species names, respectively, though this order may not always reflect the real systematic relationships. The accounts first feature the currently valid scientific name in italics, the name(s) of the first describer(s) and the year of description. If there have been changes in the genus and species name in the past, the authority is given within parentheses (brackets). They are followed by a list of synonyms (without any sign in-between the scientific name and the authority) and chresonyms (with a hyphen in-between the scientific name and the authority) given according to alphabetic order of the genus name. Subsequently the type locality (if known) and the current location the type specimen is deposited is given. Junior synonyms of currently valid taxa are given even if they have not been applied in connection with Sri Lanka. The common English- E, Sinhala- S and Tamil- T name(s) are given depending on availability. However we are aware of several cases where misidentifications were made by using vernacular names when working with local inhabitants as many people are not aware of the specific Sinhala vernacular names and the common names in general use are referred to a group of animals rather than a specific species and also local names may change with the location. Notes are included where some comment is pertinent. Unless otherwise mentioned, synonyms, chrysonyms, range and type specimen data are based on Uetz & Hallermann (1995-2008), the latter which is in turn a compilation of reliable, published sources. Distribution around Sri Lanka is based on Somaweera (2006).

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Family Hydrophiidae Boie, 1827

Astrotia Fischer, 1855 01. Astrotia stokesii (Gray, 1846)

Astrotia schizopholis - Fischer 1856 Astrotia stokesi - Wall, 1909; - Wall, 1921; - Smith 1943 Astrotia stokesii - Cogger 1983; - Cogger 2000 Disteira stokesii - McDowell 1972; - Boulenger 1896; - Rasmussen 1997; - Kharin 2005; - Grandison 1978 Hydrophis guttata Murray 1887 Hydrophis schizopholis Schmidt 1846; - Duméril & Bibron 1854 Hydrus annulatus Gray 1849 Hydrus major Shaw 1802 (part.) Hydrus stokesii Gray 1846 Global distribution: Australia, China, Gulf of Siam, India, Indonesia, New Guinea, Pakistan, Philippine Islands, Sri Lanka, Taiwan, Vietnam, West Malaysia. Records from Sri Lanka: Shallow and deep waters in North, North-west, West and South Sri Lanka. Known from Colombo, Galle, Panadura, Peysalai, Pt. Pedro, Puttalam. Type locality: Australian seas. Holotype: BMNH 1946. 1. 17. 12.

Enhydrina Gray, 1849 02. Enhydrina schistosa (Daudin, 1803)

Disteira russelii Fitzinger 1827 Disteira schistosa - Mcdowell 1972; - Grandison 1978 Enhydrina schistosa Stoliczka 1870; - Smith 1943; - Glaw & Vences 1994; - Cox et al., 1998; - Murphy, Cox & Voris 1999; Cogger 2000 Enhydrina valakadien - Boulenger 1890; Boulenger 1896; - Flower 1899 Enhydrina valakadyen - Gray 1849; - Wall 1921 Enhydrina valakadyn - Stejneger 1907 Enhydrina vikadien - Boettger 1892 Hydrophis bengalensis Gray 1842 Hydrophis fasciatus - Jan 1872 Hydrophis schistosa Schlegel 1837; - Fischer 1856 Hydrophis schistosus Daudin 1803; - Duméril & Bibron 1854; - Jan 1872 Hydrophis subfasciata Gray 1842 Hydrus valakadyn Boie 1827 Polyodontes annulatus Lesson 1834 Thalassophis werneri Schmidt 1852 Global distribution: Australia, Bangladesh, Burma, India, Madagascar, Malaysia (incl. Sabah, Sarawak), New Guinea, Oman, Pakistan, Seychelles, Sri Lanka, Thailand, United Arab Emirates, Vietnam. Records from Sri Lanka: Shallow and deep waters in North, North-west and West Sri Lanka. Known from Beruwala, Jaffna, Kalutara, Moratuwa, Negambo, Puttalam, Waskaduwa, Wilpattu. Type locality: Tranquebar, India. Holotype: BMNH 1946.1.10.7.

Hydrophis Latreille in Sonnini & Latreille, 1801 03. Hydrophis bituberculatus Peters, 1873

Aturia bituberculata - Welch 1994 Chitulia bituberculata - Kharin 2005 Distira bituberculatus - Boulenger 1896 Hydrophis bituberculatus Peters 1872; - Smith 1943; - Rasmussen 1992 Hydrophis cyanocinctus Deraniyagala 1955 (part.) Lioselasma bituberculatus - Wall 1921

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Global distribution: Sri Lanka, West coast of peninsular Thailand. Records from Sri Lanka: Capt. Nietner collected the type specimen from Colombo in 1872 (Rasmussen, 1992). No confirmed, subsequent records are found in Sri Lanka though an unconfirmed record occurs from Puttalam lagoon (CEA/Euroconsult. 1995.). The species is reported to be common around Phuket in Thailand (Rasmussen, 1992). Type locality: Colombo, Sri Lanka (as ‘Ceylon’). Holotype: ZMB 4647 (female).

04. Hydrophis cyanocinctus Daudin, 1803

Distira cyanocincta - Boulenger 1896; - Stejneger 1907; - Kharin 2005 Hydrophis aspera Gray 1849 Hydrophis chittal Rafinesque 1817 Hydrophis crassicollis Anderson 1871 Hydrophis cyanocincta - Boulenger 1887 Hydrophis cyanocinctus Daudin 1803; - Smith 1943; - Murphy, Cox & Voris 1999 Hydrophis phipsoni Murray 1887 Hydrophis striata - Schlegel 1837; - Fischer 1856 Hydrophis striatus - Duméril & Bibron 1854 Hydrophis subannulata Gray 1849 Hydrophis taprobanica Haly 1887 Hydrophis tenuicollis Peters 1873 Hydrophis trachyceps Theobald 1870 Hydrophis westermani Jan 1859 Hydrophis westermanni - Jan 1863 Leioselasma [sic] cyanocincta - Wall 1921 Leioselasma cyanocincta - Prater 1924; - Welch 1994 Leioselasma cyanocinctus - Kharin 1984; - Das 1996 Leioselasma striata Lacépède 1804 Global distribution: Bangladesh, Burma, China, India, Indonesia (Java, Aru Islands), Iran, Japan, Korea, Malaysia, New Guinea, Oman, Pakistan, Philippines, Solomon Islands, Sri Lanka, Thailand, United Arab Emirates, Vietnam. Records from Sri Lanka: Apparently found all around Sri Lanka. Known from Beruwala, Chilaw, Colombo, Dehiwala, Gogawaya, Jaffna, Kalpitiya, Kaluthara, Kandakuliya, Kumana, Mannar, Moratuwa, Negambo, Puttalam, Rekawa, Rumassala, Ussangoda, Wadduwa, Waskaduwa, Wellawatte. Type locality: “Coromandel”. Probably referring to the south east coast of the Indian Peninsula named as the Coromandel coast, the English interpretation of Chola Mandalam- the land of the Cholas. A ‘Coromandel Peninsula’ exists in New Zealand, but considering the distribution of the species the earlier place may be more related.

05. Hydrophis gracilis (Shaw, 1802)

Disteira gracilis - Fitzinger 1826; - Wall 1909 Hydophis guentheri Murray 1884 Hydrophis (Microcephalophis) gracilis - Kharin 2004 Hydrophis gracilis - Duméril, Bibron & Duméril 1854; - Fischer 1856; - Günther 1864; - Boulenger 1896; - Bauer & Vindum 1990; - Welch 1994; - Cogger 2000; - Leviton et al., 2003; - David et al., 2004; - Greer 2006 Hydrophis microcephala - Fischer 1856 Hydrophis microcephalus - Duméril, Bibron & Duméril 1854 Hydrophis rostralis Smith 1917 Hydrus gracilis Shaw 1802 Hydrus kadellnagam Boie 1827 Microcephalophis gracilis - Lesson 1834; - Wall 1921; - Smith 1943: - Das 1996; - Grossmann & Tillack 2001 Microcephalophis gracilis gracilis Smith 1926

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Microcephalophis gracilis microcephalus - Kharin 2005 Thalassophis microcephala Schmidt 1852 Global distribution: Australia, Bangladesh, Burma, China, Fuji, India, Indonesia, Iran, Melanesia, New Guinea, Oman, Pakistan, Peninsular Malaysia, Philippines, Sri Lanka, Thailand, United Arab Emirates, Vietnam. Records from Sri Lanka: Beruwala, Chilaw, Potuwil, Pt. Pedro, Trincomalee, Ussangoda. Type locality: not given (fide O’Shea 1996). Holotype: BMNH 1946.1.3.89.

06. Hydrophis lapemoides (Gray, 1849)

Aturia lapemoides Gray 1849; - Welch 1994 Chitulia lapemoides - Kharin 2005 Distira lapemoides Wall 1909 Hydrophis holdsworthii Günther 1872 Hydrophis lapemidoides [sic] Kennedy 1937 Hydrophis lapemoides Günther 1864; Smith 1926; - Smith 1943; - Rasmussen 1993 Hydrophis stewartii Anderson 1872 Lioselasma [sic] lapemidoides [sic] - Wall 1921 Global distribution: India, Iran, Malaysia (Penang island), Oman, Pakistan, Singapore, Sri Lanka, Thailand, United Arab Emirates. Records from Sri Lanka: Mainly known from shallow waters in West Sri Lanka. Known from Beruwala, Dehiwela, Jaffna, Kalutara, Kandakuliya, Moratuwa, Negambo, Waskaduwa.

07. Hydrophis mamillaris (Daudin, 1803) Anguis mamillaris Daudin 1803 Aturia mamillaris - Welch 1994 Chitulia mamillaris - Kharin 2005 Hydrophis mamillaris - Smith 1943; - Das 1996 Hydrophis mammillaris [sic] - Khan 2002 (pers. comm.) Leioselasma mamillaris - Smith 1926 Lioselasma [sic] mamillaris - Wall 1921 Global distribution: India, Pakistan, Sri Lanka. Records from Sri Lanka: A specimen collected from Pt. Pedro is lodged at the National Museum in Sri Lanka (NMSL 2005.21.1).

08. Hydrophis ornatus (Gray, 1842) Aturia ornata Gray 1842; - Welch 1994 Chitulia ornate - Kharin 2005 Distira andamanica Annandale 1905 Distira mjobergi Lönnberg & Andersson 1913 Hydrophis ellioti Günther 1864; - Boulenger 1887 Hydrophis inornatus - Smith 1926 Hydrophis laevis Lütken 1863 Hydrophis ornata - Günther 1864; - Smith 1943; - Murphy, Cox & Voris 1999; - Cogger 2000 Hydrophis ornatus - Tamiya et al., 1983; - Rasmussen 1989 Global distribution: Australia, Guangxi, Hainan, Hong Kong, India, Indonesia, Iran, Japan (Ryukyu), Malaysia, New Guinea, Oman, Pakistan, Philippines, Shandong (China), Solomon Islands, Sri Lanka, Taiwan, Thailand, United Arab Emirates, Vietnam. Records from Sri Lanka: Chilaw, Colombo, Galle, Hikkaduwa, Kalpitiya. Type locality: not given Holotype: BMNH 1946.1.23.72.

09. Hydrophis spiralis (Shaw, 1802)

Hydrophis spiralis Gray 1849; - Duméril & Bibron 1854; - Smith 1943; - Murphy, Cox & Voris 1999; - Bauer & Sadlier 2000; - David et al., 2004

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Hydrus spiralis Shaw 1802 Leiocephalus spiralis - Das 1996 Leioselasma spiralis - Prater 1924; - Kharin 1984; - Welch 1994; - Kharin 2005 Lioselasma [sic] spiralis - Wall 1921 Global distribution: China, India, Indonesia, Iran, Malaysia, New Caledonia/Loyalty Islands, New Guinea, Oman, Pakistan, Persian Gulf, Philippines, Sri Lanka, Thailand, United Arab Emirates. Records from Sri Lanka: Apparently found all around Sri Lanka. Known from Beruwala, Bundala, Chilaw, Colombo, Godawaya, Hambanthota, Kalametiya, Kalpitiya, Kalutara, Kandakuliya, Kumana, Mannar, Moratuwa, Panadura, Puttalam, Rekawa, Rumassala, Ussangoda, Waskaduwa. Type locality: Indian Ocean. Holotype: BMNH 1946.1.6.94.

10. Hydrophis stricticollis Günther, 1864

Aturia stricticollis - Welch 1994 Chitulia stricticollis - Kharin 2005 Hydrophis cf. stricticollis - Baier 2005 Hydrophis obscurus Boulenger 1896 Hydrophis stricticollis Günther 1864; - Smith 1943; - Leviton et al., 2003 Global distribution: Bangladesh, Burma, India, Sri Lanka. Records from Sri Lanka: Kandakuliya, Mannar and Puttalam in North-west Sri Lanka. Type locality: India. Holotype: BMNH 1946.1.6.90.

Kerilia Gray, 1849 11. Kerilia jerdonii (Gray, 1849)

Distira jerdonii Boulenger 1896 Kerilia jerdoni - Rasmussen & Andersen 1990; - Murphy, Cox & Voris 1999 Kerilia jerdonii Gray 1849; - Wall 1921; - Smith 1943 Global distribution: Burma, China, Gulf of Siam, India, Indonesia (Borneo), Malaysia, Mergui Archipelago, Sri Lanka, Thailand. Records from Sri Lanka: Colombo, Hambantota, Jaffna, Mannar. Type locality: Madras, India. Holotype: BMNH III.8.1.a.

Lapemis Gray, 1835 12. Lapemis curtus (Shaw, 1802)

Enhydris curtus - Boulenger 1896 Hydrophis (Pelamis) pelamidoides - Fischer 1856 Hydrophis (Pelamis) pelamidoides var. annulata Fischer 1855; Fischer 1856 Hydrophis abbreviatus Jan 1863 Hydrophis brevis Jan 1863 Hydrophis fayreriana Anderson 1871 Hydrophis hardwickei - Boettger 1888 Hydrophis hardwickii Gritis &Voris 1990 Hydrophis pelamidoides Schlegel 1837; - Duméril & Bibron 1854 Hydrophis pelamoides Hilgendorf 1876 Hydrophis propinquus Jan 1859 Hydrus curtus Shaw 1802 Lapemis curtus - Gray 1842; - Rasmussen 1997; - Murphy, Cox & Voris 1999 Lapemis loreatus Gray 1843 Global distribution: Australia, Burma, China, Fiji, India, Indonesia, Iran, Japan, Malaysia (incl. Sabah, Sarawak), New Guinea, Oman, Pakistan, Philippines, Sri Lanka, Taiwan, Thailand, United Arab Emirates, Vietnam. Records from Sri Lanka: A common species in the locations where they occur. Known from Bundala, Chilaw, Hambantota, Kalpitiya, Kumana, Panadura, Panama, Potuwil, Ussangoda.

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Type locality: East India? Holotype: BMNH 1946.1.17.59 (formerly BMNH III.2.1.a).

Pelamis Daudin, 1803 13. Pelamis platurus (Linnaeus, 1766)

Anguis platura Linnaeus 1766 Hydrophis (Pelamis) bicolor — Fischer 1856 Hydrophis (Pelamis) bicolor var. alternans Fischer 1855; Fischer 1856 Hydrophis (Pelamis) bicolor var. sinuata - Fischer 1855; - Fischer 1856 Hydrophis pelamis Schlegel 1837 Hydrus bicolor Schneider 1799 Hydrus platurus - Boulenger 1896; - Stejneger 1907 Pelamis bicolor - Schneider 1799; Daudin 1803; - Duméril, Bibron & Duméril 1854 Pelamis bicolor var. sinuata Duméril, Bibron & Duméril 1854 Pelamis bicolor var. variegata Duméril, Bibron & Duméril 1854 Pelamis ornata Gray 1842 Pelamis platura - Böhme 2003 Pelamis platuros [sic] Daudin 1803 Pelamis platurus - Smith 1943; - Peters & Orejas-Miranda 1970; - Stebbins 1985; - Liner 1994; - Glaw & Vences 1994; - Cox et al., 1998; - Murphy, Cox & Voris 1999; - Cogger 2000; - Leviton et al., 2003; - Sharma 2004 Pelamis platyura [sic] - Das & Yaakob 2007 Pelamis schneideri Rafinesque 1817 Global distribution: Probably the most widely distributed sea snake in the world. Known from Andaman Islands, Australia, China, Colombia, Costa Rica, Ecuador, El Salvador, Galapagos Islands, Guatemala, Honduras, India, Japan, Korea, Madagascar, Malaysia, Maldives, Mexico, New Caledonia, New Guinea, New Zealand, Nicaragua, Nicobar Islands, Oman, Pakistan, Panama, Peru, Philippines, Russia, Solomon Islands, Sri Lanka, Taiwan, Tanzania, Thailand, United Arab Emirates, Vietnam. Records from Sri Lanka: Batticaloa, Beruwela, Bundala, Chilaw, Dehiwela, Kalutara, Moratuwa, Negambo, Panadura, Panama, Rekawa, Rumassala, Trincomalee, Ussangoda, Wadduwa, Waskaduwa.

Praescutata Wall, 1921 14. Praescutata viperina (Schmidt, 1852)

Diseira praescutata Duméril, Bibron & Duméril 1854 Disteira viperina - Stejneger 1907 Distira viperina - Boulenger 1896 Hydrophis doliata Fischer 1856 Hydrophis jayakari Boulenger 1887 Hydrophis nigra 1872 Hydrophis plumbea Murray 1887 Hydrophis schistosus Boulenger 1896 Praescutata viperina - Wall 1921; - Smith 1943; - David & Ineich 1999; - Van Der Kooij 2001; - Khan 2002; - Sharma 2004; - Kharin2005 Thalassophina viperina - Smith 1926; - Golay et al., 1993; - Welch 1994;- Rasmussen 1997; - Murphy, Cox & Voris 1999 Thalassophis viperina Schmidt 1852; - David et al., 2004 Thalassophis viperinus - Grossmann & Tillack 2001 Global distribution: Burma, India, Indonesia (Java, Borneo, Sumatra), Malay Peninsula, Oman, Pakistan, Sri Lanka, Taiwan, Thailand, United Arab Emirates, Vietnam. Records from Sri Lanka: Galkissa, Mannar, Pt. Pedro. Type locality: Java. Holotype: ZMH 404.

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Family : Laticaudidae Cope, 1879 Laticauda Laurenti, 1768 15. Laticauda colubrina (Schneider, 1799)

Anguis platura Lacépède 1790 (fide Heatwole et al., 2005) Coluber laticaudatus Linnaeus 1758 (part.) Coluber platycaudatus Oken 1836 Hydrophis colubrina - Schlegel 1837 Hydrus colubrinus Schneider 1799; - Begbie 1846 Laticauda colubrina - Stejneger 1907; - Smith 1943; - Peters & Orejas-Miranda 1970; - Liner 1994; - Cox et al., 1998; - Cogger 2000 Laticauda scutata Cantor 1847 (not of Laurenti 1768) Platurus colubrinus - Wagler 1830; - Fischer 1884; - Fischer 1888; - Boulenger 1896 Platurus fasciatus Latreille 1801 Platurus fasciatus var. colubrina - Fischer 1856 Platurus laticaudatus var. B. - Günther 1858 (part.) Global distribution: Andaman Islands, Australia, Burma, China, Fiji, El Salvador, India, Indonesia (Borneo), Japan (Ryukyu Islands), Malaysia, Melanesia, Mexico, New Caledonia, New Guinea, New Zealand, Nicaragua, Nicobar Islands, Philippines, Polynesia, Solomon Islands, Sri Lanka, Taiwan, Thailand. Records from Sri Lanka: There are no records of this species coming ashore in Sri Lanka. Type locality: not given. Holotype: ZMB 9078.

In addition to the aforementioned ‘sea snakes’, the Wart Snake or Cloth Snake (Acrochordus granulatus) of family Acrochordidae is sometimes found in shallow sea waters and the Dog-faced Water Snake (Cerberus rynchops) and the Glossy Marsh Snake (Gerarda prevostiana) of family Colubridae live in the intertidal zones of Sri Lankan waters. Key to the sea snakes of Sri Lanka Accurate identification of sea snakes to the species level is very difficult, especially if dealing with live animals. Most species (especially Hydrophis species) show wide interspecific variation which makes it difficult to exclusively use external characters for identification (Rasmussen, 2000). Hence, this key should be used in conjunction with the existing, more-detailed larger books and scientific papers available. We have omitted scale counts as much as possible and teeth counts and bone length ratios completely. However, these features are very important in confirming a species. We have illustrated most of the features, thus in addition to the herpetologists, this key should be usable by divers, Navy personnel and medical professionals to identify a sea snake and will be most useful for live or freshly killed specimens whose colour pattern is still visible. The key is based on Rasmussen (2000) and Somaweera (2006) and all sketches are reproduced from Cogger (2000) and Rasmussen (2000).

Sea snake bites in Sri Lanka: Sea snakes are among the most venomous snakes in Sri Lanka, and according to data from other countries, most species may be more toxic than the highly venomous land snakes in the island. Most sea snake species found around Sri Lanka are gentle and inoffensive and will only bite when provoked, but certain species like Astrotia stokesii, Enhydrina schistose and Hydrophis ornatus are known to be much more aggressive (Guinea, 1994; Heatwole & Cogger, 1994; Rasmussen, 2000; Toriba, 1994; Warrell, 1994). However sea snake bites are encountered very infrequently in Sri Lanka but among them those associated with non-envenoming (‘dry bites’) are frequent. Typical victims are fishermen who try to remove sea snakes entangled in gill nets. Reid (1961) wrote that sea snake bites are usually painless and envenoming does not cause local signs. de Silva (1994) states that he could only find one reference to sea snake bite treatments among the ~150 traditional snake bite treatment literature, indicating that either the ancient physicians were unaware of their presence or that sea snake bites were never reported at that time. However, according to local beliefs, a person bitten by a sea snake should not come out of the sea and should drink sea water three times (de Silva, 1990). Thanabalasundram & Vidyasagara (1969) mentioned that sea snake bites were “not commonly seen” in Colombo. The following case reports are available with regard to Sri Lanka:

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* Polyvalent land-snake antivenom has been used in the management of these patients since specific antivenom against sea-snake envenoming is not available in Sri Lanka. Rasmussen (2000) recommends the following first-aid procedures in case of a sea snake bite: if the bite is on an arm or leg, a broad crepe bandage (or material of similar type) should be wrapped immediately around the area of the bite. The bandage must be very tight and extended over the entire arm or leg. Then a splint should be used to immobilize the arm or leg and hospital treatment must be sought as quickly as possible. If the bite is on the body, firmly press the area of the bite and look for hospital treatment immediately. Threats and conservation: Compared to terrestrial reptiles, sea snakes are not under any severe threats in Sri Lanka. Apart from direct threats like death due to entangling in fishing nets (dying from drowning or get killed by fishermen) and indirect threats including coral reef and mangrove destruction and oceanic pollution, sea snakes do not face any severe threats. They are not exploited for their skin, organs or meat in Sri Lanka, nor they are used as animal food (as in some South East Asian countries, where hundreds end up as croc food in crocodile farms). However, the latter threats persist in certain adjacent countries. Sea snakes are not listed as “protected” in the Fauna and Flora Protection Act (FFPA) of Sri Lanka and are also not protected by CITES. Acknowledgments Our sincere thanks go to Arne Redsted Rasmussen, Aaron Lobo, John Rudge, Mohomed M. Bahir and anonymous reviewers for their valuable comments.

Literature Cited Amarasekera, N., A. Jayawardena, A. Ariyaratnam, U. C. Hewage and A. de Silva, 1994. Bite of a sea snake (Hydrophis spiralis): a case report from Sri Lanka. Journal of Tropical Medicine and Hygiene, 97 (4): 195-198. Boulenger, G. A., 1896. Catalogue of the snakes in the British Museum (Natural History). Vol. 3, British Museum (Natural History), London: xiv+727. Branch, B. 1998. Field Guide to Snakes and Other Reptiles of Southern Africa. Third Revised Edition. Ralph Curtis Books Publishing, Sanibel Island, Florida. Burger, W. L. and T. Natsuno, 1974. A new genus for the Arafura smooth sea snake and redefinitions of other sea snake genera. The Snake, 6: 61-75. CEA/Euroconsult, 1995. Wetland Site Report– Puttalam lagoon, Dutch Bay and Portugal Bay. Book No. 34. Cogger, H. G. 2000. Reptiles and Amphibians of Australia. Reed New Holland, Sydney, Australia: 808 Cogger, H., H. Heatwole, Y. Ishikawa, M. McCoy, N. Tamiya and T. Teruuchi, 1987. The status and natural history of the Rennell Island Sea Krait, Laticauda crockeri (Serpentes: Laticaudidae). Journal of Herpetology, 21 (4): 255-266. Cogger, H. G. & H. Heatwole, 2006. Laticauda frontalis (de Vis, 1905) and Laticauda saintgironsi n.

Source Species involved Victim Symptoms

1 Senanayake et al., 2005 Pelamis platurus (70cm)

7-year-old boy

No local or systemic effects recorded other than a 2.5cm linear scratch mark. Hospitalised for one and a half days.

2 Senanayake et al., 2005 Enhydrina schistose (110cm)

39-year-old male

Mild redness around two bite marks but no pain or local/ systemic effects. Only prescribed tetanus toxoid.

3 Amarasekera et al., 1994 Hydrophis spiralis Pain at the bite site, regional lymph node enlargement and absence of muscle pain and tenderness. *

4 Karunaratne & Panabokke, 1972

Unidentified (Whitaker & Captain, 2004) suspects it to be Pelamis platurus

Adult fisherman

Ptosis, difficulty in talking, swallowing but no heart or respiratory difficulties. Later developed severe pains, renal failure, hyperkaemia which lasted for 24 days, and the patient died. *

5 Jahubar et al., 1984; Subramaniam & James, 1985

Reports three bites in fisherman at Mannar during a period of five months

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sp. from Vanuatu and New Caledonia (Serpentes: Elapidae:Laticaudinae) - a new lineage of sea kraits? Records of the Australian Museum, 58: 245-256. Cozzi, C. A. 1980. The Absence of Sea Snakes in the Atlantic Ocean. Bulletin Maryland Herpetological Society, 16 (3): 113-118. Das, I. and A. de Silva, 2005. A Photographic guide to snakes and other reptiles of Sri Lanka. New Holland Publishers (UK) Ltd: 144. de Silva, A. 1990. Color guide to the snakes of Sri Lanka. R & A Publishing Limited, England: vi+130. de Silva, A. 1994. An Account of the Sea snakes (Serpentes: Hydrophiida) of Sri Lanka. 234-249. In: Gopalakrishnakone, P., (Ed.) Sea Snake Toxinology, Singapore University Press: 350. de Silva, A. 2006. Current status of the reptiles of Sri Lanka. In: Bambaradeniya, C.N.B. (Ed.). Fauna of Sri Lanka: Status of taxonomy, research and conservation. IUCN Sri Lanka: 34-163. de Silva, P. H. D. H. 1980. Snake Fauna of Sri Lanka - with special reference to skull, dentition and venom in snakes. National Museums of Sri Lanka: xi, 472. Deraniyagala, P. E. P. 1955. A colored atlas of some vertebrates from Ceylon - Serpentoid Reptilia. Vol. 3, Ceylon National Museums, Colombo: 121. Dowling, H. G., 1959. Classification of the serpents: A critical review. Copeia, 1959: 38-52. Dowling, H. G., 1967. Hemipenes and other characters in colubrid classification. Herpetologica, 23: 138-142. Dunson, W. A. (Ed.). 1975. The biology of the sea snakes. London-Tokyo, University Park Press Baltimore: 530 Fernando, M., W. Gooneratne, 1983. Sea snake envenoming. Ceylon Medical Journal, 28: 131-143. Fichter G. S. 1982. Poisonous Snakes. A First Book. Franklin Watts: 66. Fry, G. C., D. A. Milton, and T. J. Wassenberg, 2001. The reproductive biology and diet of sea snake bycatch of prawn trawling in northern Australia: characteristics important for assessing the impact on populations. Pacific Conservation Biology, 7 (1): 55–73. Green, H. W. 1997. Snakes: the Evolution of Mystery in Nature. University of California Press): 366.

Guinea, M. L. 1994. Sea snakes of Fiji and Niue. 212-233. In: Gopalakrishnakone, P., (Ed.) Sea Snake Toxinology, Singapore University Press: 350. Guinea, M. L. 2003. Ecology, Systematics and Biogeography of Sea Snakes. Faculty of Education Health and Science, Northern Territory University, Darwin, Australia. Haputhantri, S. S. K., M. C. S. Villanueva and J. Moreau, 2008. Trophic interactions in the coastal ecosystem of Sri Lanka: An ECOPATH preliminary approach. Estuarine, Coastal and Shelf Science, 76 (2): 304-318. Heatwole, H., 1997. Marine snakes: are they a sustainable resource? Wildlife Society Bulletine, 25: 766-772. Heatwole, H., 1999. Sea snakes. UNSW Press, Sydney, Australia: 148. Heatwole, H. and H. Cogger, 1994. Sea snakes of Australia. 167-205. In: Gopalakrishnakone, P., (Ed.) Sea Snake Toxinology, Singapore University Press: 350. Heatwole, H., S. Busack & H. Cogger, 2005. Geographic variation in sea kraits of the Laticauda colubrina complex (Serpentes: Elapidae: Hydrophiinae: Laticaudini). Herpetological Monographs, 19: 1-136. Herre, A. W. C. T., 1942. Notes on Philippine sea-snakes. Copeia, 1942 (1): 7-9. Ineich, I., 1996. Geographic distribution- Serpentes. Hydrophis torquatus diadema. Herpetological Review, 27 (3): 154. Jahubar, M., A. Subramanium and R. F. James, 1984. An analysis of snake bite in Base Hospital Mannar. 2nd Annual Sessions of the Jaffna Medical Association: 9. Karunaratne, K. E. de S. and R. G. Panabokke, 1972. Sea snake poisoning – Case report. Journal of Tropical Medicine and Hygiene, 75 (5): 91-94. Kropach, C. 1975. The yellow-bellied sea snake, Pelamis, in the eastern Pacific. 185-213 In: Dunson, W., (Ed) The Biology of Sea Snakes. Univ. Park Press, Baltimore: xi + 530. McDowell, S. B., 1967. Aspidomorphus, a genus of New Guinea snakes of the family Elapidae, with notes on related genera. Journal of Zoology (London), 151: 497-543.

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McDowell, S. B., 1974. A catalogue of the snakes of New Guinea and the Solomons, with special reference to those on the Bernice Bishop Museum. Part I. Scolecophidae. Journal of Herpetology, 8: 1-57. Murthy, T. S. N., 1977. On sea snakes occurring in Madras waters. Journal of the Marine Biological Association of India, 19 (1): 68-72. Porter, R., Irwin, S., Irwin, T. and Rodrigues, K. 1997. Records of the marine snake species from the Hey-Embley and Mission Rivers, Far N Qld. Herpetofauna. 27 (2): 2-7. Rasmussen, A. R. 1989. An analysis of Hydrophis ornatus (Gray), Hydrophis lamberti (Smith) and Hydrophis inornatus (Gray) (Hydrophiidae, Serpentes) based on samples from various localities, with remarks on feeding and breeding biology of Hydrophis ornatus. Amphibia-Reptilia, 10: 397-417. Rasmussen, A. R., 1992. Rediscovery and redescription of Hydrophis bituberculatus Peters, 1872 (Serpentes: Hydrophiidae). Herpetologica, 48 (1): 85-97. Rasmussen, A. R. 1993. The status of the Persian Gulf Sea Snake Hydrophis lapemoides (Serpentes, Hydrophiidae). Bulletin of the British Museum Natural History (Zoology), (59): 97-104. Rasmussen, A. R., 1997. Systematics of sea snakes: a critical review. Symposium of the Zoological Society of London, 70: 15-30. Rasmussen, A. R., 2000. Sea snakes. At Food and Agriculture Organization of the United Nations website. Accessed 20 September 2008: 3987-4008. Rasmussen, A. R., M. Auliya and W. Bohme, 2001. A new species of sea snake genus Hydrophis (Serpentes: Elapidae) from a river in West Kalimantan (Indonesia, Borneo). Herpetologica, 57 (1): 23-32. Reid, H. A., 1961. Diagnosis prognosis and treatment of sea-snake bite. Lancet, 2: 399-402. Senanayake, M. P., C. A. Ariaratnam, S. Abeywickrema and A. Belligaswatte, 2005. Two Sri Lankan cases of identified sea snake bites, without envenoming. Toxicon, 45: 861-863. Smith, M., 1926. Monograph of the Sea-snakes (Hydrophiidae). London, British Museum (Natural History): 130.

Somaweera, R., 2006. Sri Lankawe Sarpayin (‘The Snakes of Sri Lanka’). Wildlife Heritage Trust of Sri Lanka: 297. Thanabalasundrum, R. S. and N. W. Vidyasagara, 1969. Snake bites and its treatment. Ceylon Medical Journal, 14 (4): 188-191. Toriba, M., 1994. Sea snakes of Japan. 206-211. In: Gopalakrishnakone, P. (Ed.) Sea Snake Toxinology, Singapore University Press: 350. Uetz, P. and J. Hallermann, 1995-2008. The TIGR Reptile Database. Retrieved on 20 April 2008, from http://www.reptile-database.org/. Underwood, G., 1967. A contribution to the Classification of Snakes. London: Trustee British Museum (Natural History). Underwood, G., 1979. Classification and distribution of venomous snakes in the world. In: Lee, C. Y. (Ed.). Handbook of Experimental Phamacology. Vol. 52. Snake Venoms. Berlin: Springer-Verlag: 16-40. Voris, H. K., H. H. Voris, and L. B. Liat, 1978. The food and feeding behavior of a marine snake, Enhydrina schistosa (Hydrophiidae). Copeia. 1978: 134-146. Voris, H. K. and B. C. Jayne, 1979. Growth, reproduction and population structure of a marine snake, Enhydrina schistose (Hydrophiidae). Copeia. 1979 (2): 307–318. Wall, F., 1909. A monograph of the sea snakes (Hydrophinae). Asiatic Society, Culcutta. Memoirs, 2 (8): 169-251. Wall, F., 1921. Ophidia Taprobanica or the Snakes of Ceylon. Governmental Press, Ceylon: xxii+581. Warrel, D. A., 1994. Sea snake bites in the Asia-Pacific region. 1-36. In: Gopalakrishnakone, P. (Ed.). Sea Snake Toxinology, Singapore University Press: 350. Westhoff, G., B. G. Fry, H. Bleckmann, 2005. Sea snakes (Lapemis curtus) are sensitive to low-amplitude water motions. Zoology, 108: 195-200. Whitaker, R. and A. Captain, 2004. Snakes of India, Draco Books, India: 500. Zimmerman, K. and H. Heatwole, 1990. Cutaneous Photoreception: A New Sensory Mechanism for Reptiles. Copeia, 1990 (3): 860-862.

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GROVES & JAYAWARDENE, 2009


THE WILD BUFFALO OF SRI LANKA

Submitted: 20 February 2009, Accepted: 26 February 2009

Colin Groves 1 and Jayantha Jayawardene 2

1 School of Archaeology & Anthropology, Australian National University, Canberra, ACT 0200, Australia 2 Biodiversity and Elephant Conservation Trust, 615/32, Rajagiriya Gardens, Rajagiriya, Sri Lanka.

1 Corresponding author: [email protected] Abstract Buffaloes live wild in many areas in Sri Lanka, but it is controversial whether any of them are genuinely wild, or whether they are all feral. We investigated this question by observation of living buffaloes in Yala National Park, and measurement of available cranial material. We conclude that there is indeed an indigenous wild buffalo in Sri Lanka. Steps should be taken to ensure its genetic integrity. Key Words: Buffalo, Bubalus arnee migona, Sri Lanka, Yala, cranial capacity. Introduction Does Sri Lanka have a native wild buffalo? 19th-century authors, such as Kelaart, listed the Sri Lankan wild buffalo as a genuinely wild animal, and Deraniyagala (1952, amplified in 1953) described it as a separate subspecies. More recent authors, however, have either assumed it to be of feral origin (Ellerman & Morrison-Scott, 1951; Phillips, 1980), or have suspended judgment. Eisenberg & McKay (1970) wrote “status of feral population is debatable”; while Ashby & Santiapillai (1983) remarked, “Given the distinctiveness of the free-living form in Sri Lanka, the balance of probability is that it should be

regarded as wild rather than feral but no conclusive answer can be given on the present evidence”. Wherever there are wild buffalo, there is a danger of interbreeding with local domestic stock. The truly wild buffalo of the Asian mainland are all notably larger than any local domestic buffaloes, and it would not be expected that feral or mixed buffaloes would offer them much competition; consequently, feral buffaloes would have little chance of surviving for long, and gene-flow would be almost entirely one-way (from wild to domestic). If on the other hand the genuine wild stock becomes numerically depleted, competition would be relaxed, and there

TAPROBANICA, ISSN 1800-427X. April, 2009. Vol. 01, No. 01: pp. 55-62, pls 8. © Taprobanica Nature Conservation Society, 146, Kendalanda, Homagama, Sri Lanka.

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would be an opportunity for feral populations to become established with the resulting influx of domestic genes into the remnant wild population. Under such conditions, it might be difficult to disentangle unmixed from mixed stock within a wild population.

This problem was faced by Heinen (2002) in the case of the buffalo of Kosi Tappu Reserve, in Nepal. The domestic stock in the surrounding area are so-called River Buffaloes, which with their solid black colour and small, spirally curled horns are strikingly different from the wild buffaloes in the Reserve. But in the past the wild buffaloes have been persecuted, and rather few remain, so there is a problem of sorting out the mixtures. Heinen (2002) gave criteria for differentiating wild from backcrosses; among these are behavioural differences – wild herds are cohesive and consist of a single bull with a small group of cows and calves, while backcrossed herds are much more variable, and often lack bulls. Later genetic analysis (Flamand et al., 2003) seems to have corroborated Heinen’s conclusions. There could well be a similar problem in Sri Lanka, where a rinderpest epidemic in the early 20th century almost wiped out the wild buffaloes (Phillips, 1980). Whether they support the idea of a truly wild stock or not, all recent authors have agreed that there is a problem of feral and backcross stock. For example Deraniyagala (1955) acknowledged that by the time of his writing most of the wild herds had become intermixed: “The relatively purest herds are restricted to Yala Game sanctuary, but much vigilance will be necessary if this remnant is to be kept free from domestic animals which are now encroaching upon this once inaccessible area”. The domestic buffaloes of Sri Lanka are reputed to be Swamp Buffaloes; this domestic type, otherwise restricted to Southeast and East Asia, is very different in appearance from the River Buffalo breeds of South Asia, being grey or grey-brown, marked with white on the lower legs, throat and face, and with crescentic horns more resembling the wild buffalo. If this is the case, then interbreeding would be much more difficult to detect than with the River Buffaloes of Nepal. River and Swamp buffaloes have different chromosome numbers: 2n = 50 in River but 48 in Swamp buffaloes; in fact Sri Lankan buffaloes, despite their external resemblance to Swamp buffaloes, have a diploid number of 50 like River breeds (Bongso et al., 1977).

The purpose of this study is to try to determine whether there is indeed a genuine wild buffalo in Sri Lanka and, if so, what relationship it bears to those of the mainland of Asia as revised by Groves (1996). Material and methods We studied skulls of wild and domestic buffaloes in several collections; most of these are listed in Groves (1996), but some additional material was studied in the Sri Lanka National Museum (SLNM) and in the private collection of Kelum Manamendra-Arachchi (KM-A). Measurements taken are as given in Groves (1996) and in addition cranial capacities were measured where possible. The available skulls of Sri Lankan wild buffaloes were as follows (the numbers are the sequential numbers in the analysis): (1) Locality “Ceylon”: 79. CPH 1188; 83. SLNM 78G; 85. NMW 5300; 88. ZMB 32130. (2) Locality Yala: 80. SLNM 78; 86. BM 46.210; 87. FIR 8072; 92. Yala – mounted in museum of Yala Vistors’ Centre; 94. Yala – brought in to Yala Vistors’ Centre by tracker on Saturday, Aug.28, 2004; 95. Yala – brought in to Yala Vistors’ Centre by tracker on Sunday, Aug. 29, 2004. (3) Locality Gammaduwa, Knuckles Range: 89. Private collection of KM-A. These skulls were compared, using multivariate analysis (SPSS version 12.0.1), with both wild skulls from the Asian mainland and with skulls of domestic buffaloes, including both River Buffaloes (from South Asia and Italy) and Swamp Buffaloes (from Southeast Asia). If any Sri Lankan skulls fell outside the range of the domestic skulls, this would indicate “something different”, hence might be taken as evidence for the existence of a truly wild strain in Sri Lanka. Colin Groves measured the cranial capacities of as many skulls, wild and domestic, as possible. Buffalo skulls, being large, are easily damaged; the skull of a wild buffalo would often in the past be treated as a symbol of a victory of a “brave hunter” over a dangerous adversary, and mounted on a trophy shield with its base sawn off. Even skulls not so treated are often peppered with bullet holes.

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Under such circumstances, the sample available for cranial capacity determination was small, but still worth documentation. Theory dictates that domestic representatives should have, on average, a small cranial capacity, after the absolute size of the specimen has been taken into account, compared to the wild ones (Hemmer, 1983; Herre & Röhrs, 1990). Size-standardisation was by plotting cranial capacity (or endocranial volume: ECV) against the distance from basion to hormion, representing approximately the length of the brain-stem. As a final step, those Sri Lankan skulls deemed to be those of truly wild buffaloes were compared with those of Asian mainland buffaloes. We planned a visit to Yala National Park; in the event, J. Jayawardene was unable to participate but organized a visit by C. Groves in company with Mohomed M. Bahir, where as many groups of wild living buffaloes as possible were observed. Most of these were photographed, and all were briefly described and compared with the domestic buffaloes seen in the vicinity of the Park. A total of 13 groups of wild buffaloes were seen, mostly in and around waterholes in Block One of the Park. Results: wild or feral? Craniometry: In the Discriminant Analysis using horn measurements (Pl. 10; Fig. 1), the Sri Lankan sample fall partly within the domestic range, partly beyond it. Skull 89 falls within the domestic cluster and nos. 79 and 95 on the borders of it, but nos. 80, 85, 86, 88, 92 and 94 fall outside it. These, consequently, seem to be good candidates for consideration as genuine wild buffaloes. DF1 depends partly on size, but tip-to-tip distance does not follow the size trend; DF2 relies largely on tip-to-tip, base-to-tip and total span. In the analysis using cranial measurements (Pl. 10; Fig. 2), those falling within the domestic range are nos. 80, 88 and 92; those falling outside it, or on the edge, are nos. 85, 86, 90 and 94. It is interesting that, although the two datasets are quite distinct except for biorbital breadth, there is considerable agreement between the analyses as to the placement of particular specimens. DF1 relies on a contrast between greatest and least occipital breadth, and on nasal length and nasal breadth. Pl. 11; Fig. 3 depicts the difference in horn shape between domestic swamp buffaloes, to which Sri Lankan domestic buffaloes bear most resemblance (but see below), and Sri Lankan wild buffaloes. Although some wild specimens fall within the domestic range, most

do not. Note that even very large swamp buffaloes do not resemble Sri Lankan wild buffaloes, but continue the general domestic trend. One of the two domestic buffaloes with very low tip-to-tip values is a carabao from the Philippines; the other is a Sri Lankan domestic individual. Fig. 4 shows that, on the contrary, Sri Lankan wild buffaloes agree with swamp buffaloes in having a small base-to-tip distance (Pl. 11; Fig. 4a) and smaller span (Pl. 12; Fig. 4b) compared to wild samples from the mainland. Pl. 12; Fig. 5 illustrates what is perhaps the most outstanding cranial difference between wild and domestic (swamp) buffaloes: the reduced nasal length of the latter. The Sri Lankan wild sample has long nasals, like Thailand Bubalus arnee theerapati. Pl. 13; Fig. 6 shows the reduction in cranial capacity of domestic buffaloes compared to mainland wild specimens. The skull with the highest cranial capacity – and so most likely wild – Sri Lanka skull is Yala, no.94; followed by KM-A’s from the Knuckles Range; the two lowest are Yala 95 and SLNM, from Yala. Observations in and around Yala: Domestic buffaloes in Sri Lanka are reputed to resemble swamp buffaloes phenotypically (Cockrill, 1974). Their diploid chromosome number, however, is 2n=50 like those of river buffaloes (Bongso et al., 1977) and they assort with river buffaloes in the DNA sequences of 21 microsatellites and 25 polymorphic protein-coding loci (Barker et al., 1997). The domestic buffaloes of southern Sri Lanka, including the outskirts of Yala National Park (Pl. 17; Fig. 14) itself, were not in fact like any swamp buffaloes seen by CPG elsewhere (in China, Vietnam, Thailand and Indonesia). They appeared narrower in the body and shorter-legged than any swamp buffaloes, which are noticeably heavy-bodied and short-legged; in their build they were much more like river buffaloes. The horns were not flat and crescentic like swamp buffaloes, but had a distinct initial downward sweep followed by an upward curl in the distal portion especially toward the tip, not as tightly curled as in more specialised river breeds (Murrah, Italian), but very like less specialized breeds such as Pandarpuri. Colour varied from dark grey to black; swamp buffaloes are mostly dark grey, while river breeds are black. The colour pattern, however, did more resemble swamp buffaloes. Most individuals had light yellowish grey distal limb segments, grey-white muzzle and a white spot at the inner corner of each eye; most breeds of river buffaloes are unvarying black, but

57



most swamp buffaloes are dark grey with prominently grey-white distal limb segments and white marks on the face and throat. A feature that we have not seen in any domestic buffaloes elsewhere was a thick mat of medium brown hair, contrasting with the general body colour, on the forehead between the horns, and extending back as a short crest along the nape to the withers where it would form a tuft. Wild buffaloes are very numerous in Yala Block 1; in the heat of the day, every water-hole is full of them. Most of those seen were very like the domestic ones, and so undoubtedly feral; in fact, a ranger affirmed that actual domestic animals often move into the Park and spend the day soaking, but move back to the salt-licks near the villages in the evening. As well as these clearly feral or actual domestic buffaloes, however, there was a rarer type that was quite distinctive: noticeably larger and

seemingly stockier, with horns in a single plane; brownish black, with no white markings except inconspicuously along the upper lip, and lacking the brown forehead-to-withers hairs. No difference was detectable in belly or tail hair, but this could be mainly because so many of the animals seen were partly submerged. The horns had no initial downward trend, nor upward curl along their length; instead, they were in a single plane, describing an almost flat semicircle, the tips (if long enough) curving inward. All of the mature bulls that could be fully observed (Pl. 17; Fig. 15), and some cows, were of this type, which we take to be the true wild-type. Some cows appeared intermediate between the two types. The difference between wild-type horns and those apparently of mixed ancestry can be seen by comparing the two cows in Pl. 17; Fig. 16. Complete counts at seven different waterholes gave groupings as follows:

Table 01: Groups of buffalo seen in Yala National Park, Block 1 (August, 2004) Group no. No. of

Individuals Bulls Cows Calves Other notes

1 04 01: wild-type

02: one was wild-type; other was intermediate with horns bent downward

01 -

2 05 01: wild-type 04: wild-type - -

3 08 -

06: all with very long horns, rather down swept in three cases

01: accompanying one of the cows

Other animal was almost submerged, but seemed to be a bull judging by the thickness of the horns

4 09 01: very large, wild-type, black

06: grey, two with very light legs and one with extremely long, spreading horns

02: half-grown -

5 14

05: young, with short horns, one having especially white legs

09: domestic-type -

Other animals included a zebu cow at this waterhole, not far inside the Park

6 28

01: mostly submerged and could not be clearly seen

17 10 -

7 55

02: one was sub adult; other was an adult, which could not be clearly seen

23: mostly with down and upwardly curled horns, in one case asymmetrical

30: different ages, the younger ones very light colored

-

58



8 04 - -

05: two were young; three were half-grown, seen in the bush away from any waterhole; light grey, and all individuals had very light legs

-

9

A large group of cows and calves, count

was not possible

- 10< of domestic-type, were on one side of a waterhole (some are shown in Pl. 17; Fig. 17)

Well apart from them, on the far bank, was a group of three – a bull and a very large cow, both of wild-type, and another cow also very large but with some horn curl and a trace of the brown nape crest (Pl. 17; Fig. 16).

Discussion: wild or feral? The craniometric results indicate strongly that some Sri Lankan specimens represent fully or nearly pure-bred wild buffaloes, in that they fall outside the range of variation of domestic specimens. Note that both swamp and river buffaloes are included in

the domestic skull sample, but only swamp in the horn sample. Taking the results of the discriminant analyses on both skulls and horns, backed up by the bivariate scatterplots, the specimens from Sri Lanka rate as follows:

Table 02: Allocation of Sri Lankan wild-collected skulls (DR: Domestic Range)

Locality Catalog No.

“Ceylon”

79. CPH 1188 at border of DR 83. NM 78G at or beyond border of DR 85. NMW 5300 definitely outside DR 88. ZMB 32130 definitely outside DR

Yala

80. NM 78 at or beyond border of DR 86. BM 46.210 definitely outside DR 87. FIR 8072 definitely outside DR 92. Yala – mounted in museum at border of DR 94. Yala – brought in Sunday, Aug. 29 2004 definitely outside DR 95. Yala – brought in Sunday, Aug. 29 2004 At border of DR

Gammaduwa (Knuckles) 89. K.M-A within DR 90. K.M-A within DR

For safety’s sake, therefore, only the following specimens of the Sri Lankan sample will be treated as pure-bred wild buffaloes: 85. NMW 5300; 86. BM 46.210; 87. FIR 8072; 88. ZMB 32130; 94. Yala – Sunday, Aug. 29 2005. Others may also be “pure-bred” but, because their position was equivocal in one or more analyses, they will be treated cautiously, and excluded from the comparisons with mainland wild buffaloes.

Kelaart said that “the wild buffalo is of a darker colour and more hairy than the domesticated variety” (Kelaart 1852). Deraniyagala (1952), in the course of describing the Sri Lankan wild buffalo as a subspecies separate from that of India, listed some differences between it and the domestic buffaloes of Sri Lanka, as follows: “The wild Ceylon form is larger and stands higher in the leg, with reddish brown hair. Its horns are stronger, longer and more erect, with their tips tending more

59



forwards than in the domestic form... The neck of the adult male is also more developed” (Deraniyagala 1952). It will be seen that, once the obvious feral and backcrossed animals are excluded, a distinctive and homogeneous morphology does remain (see photo in Buchholtz, 1988:372-3). As described by Kelaart (1852) and Deraniyagala (1952), animals of this description, i.e. that are outside the phenotypic range of the domestic ones, are indeed larger and darker, but they cannot be described as reddish brown, nor as more hairy – note however that Kelaart was resident at Nuwara Eliya, and wild buffaloes at this altitude might indeed be (or have been) more hairy. The horns of the wild stock are flatly crescentic (in a single plane), but not necessarily more erect, unless by this Deraniyagala (1952) meant to contrast the shape with the initial downsweep of most domestic animals; if the tips tend more forwards, it is because they are generally longer, and so describe more of a semicircle. These conclusions are similar to those of Heinen (2002), who had much the same problem of distinguishing wild from feral and backcrossed buffaloes in Nepal; Nepalese domestic buffaloes, however, are of more strongly specialized river type, so their differences from wild ones are more clear-cut – for all that Sri Lankan domestics are of river stock, not swamp, they are primitive, which is doubtless why Cockrill (1972) deemed them to be swamp buffaloes. Heinen’s criteria were tested by Flamand et al. (2003), using several genetic loci; they were able to allocate wild vs. feral and backcrossed animals with a high degree of success. Heinen (2002) makes the point that wild herds have a definite social structure, of cows and calves with a herd bull, the members of the herd being always in close proximity; while feral/backcrossed groups were very variable in size and composition, and were often without herd bulls. The same features seem to distinguish wild from feral/backcrossed groups around the waterholes of Yala. We interpret the groups described earlier as follows: group 2: evidently a pure-bred wild herd; group 1: a wild bull with one wild cow, one backcrossed cow; group 4: a wild bull with feral cows; groups 3, 6, 7: the bull unclear, the cows largely backcrossed or feral; group 5: an entirely feral grouping. The “final group”, mentioned in the right-hand column of group 9, appears to be a social herd of 3 pure-bred (one bull, two cows), keeping separate from a feral/backcrossed conglomerate.

Results: wild buffalo from Sri Lanka vs. mainland: How does the Sri Lankan wild buffalo relate to the three mainland subspecies described by Groves (1996)? Is it a fourth subspecies of Bubalus arnee, or should it be ranked as a distinct species? It is also presumably possible, though not at all likely, that it represents one of the mainland subspecies, introduced in the distant past to Sri Lanka. Deraniyagala (1952) briefly gave the name Bubalus bubalis migona to the Sri Lankan wild buffalo, with the type a mounted head No. 78 B, in the Colombo museum. For this specimen (not found in my visits of 2003 and 2004) the museum catalogue gives a tip-to-tip distance of 2 feet, 7 inches (789.5mm), which is within the range of specimens measured by CPG. He gave the type locality of his new subspecies as Yala; the catalogue however lists no locality for the specimen. In the following year Deraniyagala (1953) compared to his new subspecies more specifically with the Indian wild buffalo: the horns, he stated, are relatively shorter and curve more forward towards their tips. In what follows, the name migona will applied to the Sri Lankan wild buffalo without prejudice to what its ultimate taxonomic status may be. The horns of wild buffaloes can be sexed by the basal diameter of the sheaths. As Pl. 13; Fig. 7 shows, the degree of sexual dimorphism is about equivalent in all four putative subspecies (note that CPG has reallocated the sex of some specimens of B. a. theerapati from Groves, 1996). To judge from those in specimens which both sheath and core could be measured, the core diameter is about 10 mm less than that of the sheath. Not many domestic specimens in the sample had the sexes recorded, but no bimodality in basal diameter is apparent, so it may be that this is one further character in which domestic buffaloes differs from wild. The horns of Sri Lankan buffaloes average smaller in all dimensions than those from the mainland; the distance from base to tip is shown in Pl. 14; Fig. 8, and horn span in Pl. 14; Fig. 9. The skulls of both migona and theerapati are noticeably smaller than those of arnee and fulvus, and unlike the latter the sexes are the same size (Pl. 15; Fig. 10). In the case of the greatest occipital breadth, however, migona is smaller than theerapati, and unique in its lack of sexual dimorphism (Pl. 15; Fig. 11). The narrower occiput may relate to the relatively small horns of migona.

60



Pl. 16; Fig. 12 shows a discriminant analysis of three cranial measurements (one of the five available skulls lacks the other measurements). In this admittedly limited comparison, migona overlaps with both arnee and theerapati, but is overall closer to the latter. In Pl. 16; Fig. 13, four horn measurements are used; here, the separation of migona is somewhat greater, only one of the specimens being within the ranges of the mainland samples. We may conclude that, on the data so far available, the skulls and horns of migona differ on average from those of mainland Bubalus arnee, but not absolutely. It may be different in the case of external characters. Among the ways in which the wild buffalo of Yala differ from domestic buffalo is the virtual absence of white markings. All photographs of mainland B. arnee show conspicuous grayish or yellowish white limbs, from the knees and hocks to the hoofs, white gorgets (one or two transverse crescents on the throat), and white spots on the lower face. It may be, therefore, that in this respect the Sri Lankan form is absolutely different from the mainland ones. If this is really the case, then it would be appropriate to recognize it as a distinct species (under the so-called Phylogenetic Species Concept, a species has fixed heritable differences from others). For the moment, caution should be observed, for two reasons: (1) we have closely examined wild buffaloes only in Yala: those in other areas in Sri Lanka, if they still exist, may differ; (2) On the mainland, while several wild populations of B. a. fulvus (in Kaziranga, Manas and elsewhere in Assam) are well-known, and are consistent in their possession of white markings, much less is known of the other two subspecies, although limited evidence does suggest that they too are white-marked. Until the evidence is more complete, therefore, we should refer to the Sri Lankan wild buffalo as Bubalus arnee migona, while bearing in mind that it may ultimately have to be upgraded to species rank. Postscript: “About seventy years ago”, wrote Deraniyagala (1953), “wild buffaloes abounded in all forests of the low country, but today most of them have interbred with domestic stock. The relatively purest herds are restricted to Yala Game Sanctuary, but much vigilance will be necessary of this remnant is to be kept free from domestic animals which are now encroaching upon this once inaccessible area” (Deraniyagala 1953:27). This assessment remains true today (Buchholtz, 1988).

It is perhaps not completely unexpected that there should be still some pure-bred wild stock remaining, because a feral bull would have little or no chance of dominating a wild one, and would perhaps even be dominated by a wild cow; one would then expect, on theoretical grounds, that even hybrid cows would mate only with wild bulls, so that each backcrossed generation would be graded-up. Nonetheless, action should be taken to prevent more domestic stock from entering the National Park and joining those already there; and to somehow get rid of the feral and backcrossed animals now living permanently wild. Considering that there is a widespread appreciation of the value of wild “blood” in domestic stock (for example, Heinen, 2002), obviously backcrossed animals could be sold off to local herders. There is still a good chance of preserving Bubalus arnee migona in a relatively pure state, as part of Sri Lanka’s wild heritage. Literature cited Ashby, K. R. and C. Santiapillai, 1983. The ecology of free-living water buffalo (Bubalus bubalis L.) in Sri Lanka and with particular reference to Ruhuna National Park. Tigerpaper, 10: 20-26. Barker, J. S. F., S. S.Moore, D. J. S. Hetzel, D. Evans, S. G. Tan and K. Byrne, 1997. Genetic diversity of Asian water buffalo (Bubalus bubalis): microsatellite variation and a comparison with protein-coding loci. Animal Genetics, 28: 103-115. Bongso, T. A., W. L. J. S. Kumaratileke and V. Buvanendran. 1977. The karyotype of the indigenous buffalo (Bubalus bubalis) of Sri Lanka. Ceylon Veterinary Journal, 25: 9-11. Buchholtz, C., 1988. Rinder. Grzimeks Tierleben: Säugetiere, 5: 360-417. Munich: Kindler Verlag. Cockrill, W. R., 1974. The buffaloes of Sri Lanka. In: W. R.Cockrill (Ed.). The Husbrandry and Health of the Domestic Buffalo. Rome: Food and Agriculture Organization of the United Nations. 629-635 Deraniyagala, P. E. P., 1952. Some Sinhala combative, field and aquatic sports and games. Spolia Zeylanica, 25: 179- 215. Deraniyagala, P. E. P., 1953. The wild buffalo of Ceylon, a new subspecies. Spolia Zeylanica, 27: 103-105.

61



Eisenberg, J. F. and G. M. McKay, 1970. An annoted checklist of the recent mammals of Ceylon with keys to the species. Ceylon Journal of Science, 8: 69-99. Ellerman, J. R. and T. C. S. Morrison-Scott, 1951. Checklist of Palaearctic and Indian mammals. London: Trustees of the British Museum. 383. Flamand, J. R. B., D. Vankan, K. P. Gairhe, H. Duong and J. S. F. Barker, 2003. Genetic identification of wild Asian water buffalo in Nepal. Animal Conservation, 6: 265-270. Groves, C. P., 1996. The taxonomy of the Asian wild buffalo from the Asian mainland. Zeitschrift für Saugetierkunde, 61: 327-338. Heinen, J. T., 2002. Phenotypic and behavioural characteristics used to identify wild buffalo Bubalus bubalis from feral backcrosses in Nepal. Journal of Bombay Natural History Society, 99: 173-183. Hemmer, H., 1983 (trans.1990). Domestication: the Decline of Environmental Appreciation. Cambridge: Cambridge University Press. Herre, W. and M. Rőhrs, 1990. Haustiere – Zoologisch Gesehen. 2nd edition. Stuttgart: Gustav Fischer. Kelaart, E. F., 1852. Prodromus Faunae Zeylanicae. R. Pethiyagoda and K. Manamendra-Arachchi (Eds.). Wildlife Heritage Trust, Colombo, 342. Phillips, W. W. A., 1980. Manual of the mammals of Sri Lanka. 2nd.ed. Wildlife and Nature Protection Society of Sri Lanka, Colombo.

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WALIKANNA AND KARUNARATHNA, 2009


NOTES ON EX-SITU INCUBATION AND HATCHLINGS OF Eutropis carinata (SCHNEIDER, 1801) (REPTILIA: SCINCIDAE) FROM SRI LANKA


Thilina P. Walikanna 1 and D. M. S. Suranjan Karunarathna 2 1 The Young Zoologists’ Association, Department of National Zoological Gardens, Dehiwala, Sri Lanka

2 IUCN - Sri Lanka Country office, No: 53, Horton place, Colombo 07, Sri Lanka 2 Corresponding author: [email protected] Abstract Eutropis carinata is widely distributed skink in Sri Lanka. This observation describes ex-situ incubation and hatching of this species at Colombo district in Sri Lanka. In total, 25 eggs with two adults of E. carinata were located from a termite mound while some villagers where trying to kill a cobra (Naja naja) living in the mound. Of these skink eggs, 20 were successfully incubated. The incubation period was 30 days from the date eggs were collected. The average incubation temperature varied from 26–28°C. Mean egg length was 15.6 mm and mean egg width 10.4 mm. Mean hatchling SVL was 28.5 mm and mean TL is 41.3 mm. The mean weight of hatchlings was ~3 g and the mean total length is 69.8 mm. After about five days, hatchlings were released at the original place of the clutch. Key Words: Eutropis carinata, ex-situ incubation, egg hatching, conservation, Sri Lanka Introduction The genus Eutropis in Sri Lanka consists of seven species: E. beddomei, E. bibronii, E. carinata, E. floweri, E. macularia, E. madaraszi and E. tammanna (Deraniyagala, 1953; Das & de Silva, 2005; Das et al., 2008). The Common Skink, Eutropis carinata (Schneider, 1801) is known locally in Sinhala as “Sulaba Garandi Hikanala”. This species is the largest skink in Sri Lanka (de Silva, 1996, 2006). According to the published

literature, E. carinata has been widely distributed in Sri Lanka. The habitats are characterized as open areas, forests with close canopies, home gardens as well as plantations in wet and dry zones below 1,000 m a.s.l. (Das & de Silva, 2005). Adults of E. carinata measured from snout to vent of 140< mm, a head length of 25< mm, a tail length of 200< mm and an axilla to groin length of 100< mm (Deraniyagala, 1953). The present communication


63

OBSERVATIONAL NOTES ON EX-SITU INCUBATION AND HATCHLINGS OF Eutropis carinata


adds to our current knowledge on ex-situ incubation and hatching of E. carinata. Methods of incubation: An enclosure is built of 5 mm thick, waterproof glass and these form the base, the front, the back and both sides. The top consists of extruded plastic fly screen framing and this allows free flow of air but can be covered with plastic sheeting to increase humidity if desired. The lid is close fitting to protect the eggs from predators. The length of the enclosure is 30 cm, width 15 cm and height 10 cm. A thermometer and a hygrometer are used to monitor temperature and relative humidity fluctuations. The base of the unit is filled with soil mixed with sand to a depth of nearly 4 cm. A few pieces of stones and leaf litter were also provided as places of concealment for hatchlings. The relative humidity varied between about 60–74% during the month of incubation. These values may be much higher than those found in the open in the wild. The surface soil was normally kept dry but occasionally about 30 ml of water was sprayed in to the hatching device to simulate more natural conditions. The temperature range was 26–28°C. The daily variation varied about 1–2°C. The 25 incubating eggs were half-buried in the soil and covered with leaf litter. After about five days, the hatchlings were released to the original habitat. Observations On the 30 January 2008 we found 25 eggs, inside a termite mound at 10:30 hr. The eggs were incubated in Colombo district in Sri Lanka. Of the 25 eggs, 20 were successfully incubated. The incubation period was 30 days from the date eggs were captured, at an average incubation temperature of around 27°C (range 26–28°C). The mean length of the eggs was 15.6 mm (range 13.7–18.0 mm) and mean width was 10.4 mm (range 9.2–14.8 mm) (Fig. 1). Fig. 01: Three of 20 hatched E. carinata eggs

Measurements of the eggs and hatchling data are presented in the Table 01. The mean SVL of the hatchlings was 28.5 mm (range 26.2–30.2 mm) and mean TL was 41.3 mm (range 36.5–46.3 mm) (Fig. 2). The mean weight of the hatchlings was ~3 g (based on 10 specimens) (range 4–6 g) and the mean total length was 69.8 mm (range 62.7–76.5 mm). Fig. 02: Newly hatched E. carinata Discussion We have located 25 eggs with two adults (the male and the female) while some villagers were trying to kill a cobra (Naja naja) living in a termite mound. The cobra was about 1.5 m total length and the two skinks and eggs were present in this termite mound. The interesting fact about this observation is that this is probably the first record of E. carinata from a termite mound which inhabited by a cobra. This is the first documented hatching of E. carinata eggs observed from Sri Lanka. Sometimes E. carinata and N. naja may show commensalism; however, there are no previous records of this kind of behaviour for E. carinata. Somehow, it is a better habitat for E. carinata to live with a N. naja in a termite hole, because we assume that other snakes, especially Ptyas mucosa, Cercaspis carinata, Olidodon arnensis, Lycodon aulicus, Boiga ceylonensis, Hypnale hypnale and Dendelaphis tristris, never reach there because the ophiophagous Naja naja is one of their natural predators. In addition, Varanus bengalensis and some birds like White breasted King fisher (Halcyon smyrnensis), Gray horn bill (Ocyceros gingalensis), Shikra (Accipiter badius) and Little Scops Owl (Otus bakkamoena) are natural predators of E. carinata. It is not clear for us why Naja naja refrain from feeding on E. carinata adults living inside the same termite hole. We have observed Naja naja feed on toads, other snakes, Naja naja, juveniles of Varanus bengalensis as well as agamid lizards, but we never observed Naja naja feeds on skinks before, but skinks were swallowed by cobra as well as Ceylon

64

WALIKANNA AND KARUNARATHNA, 2009


krait, Bungarus ceylonicus (A. de Silva, November 2008, pers. comm.). Sometimes the skink may use another root to enter the termite mound where those eggs were deposited. However we could not examine the entire hole because while we reach there the villagers had broken a half of the termite mound. According to the perused literature information on behaviors, feeding and breeding habits, population dynamics, ecology and threats are not properly understood for most species in Sri Lanka (Amarasinghe, 2009). Therefore, we believe that this information on ex-situ incubation and hatching of E. carinata may facilitate the future conservation of this species as well as other skinks. Table 01. Measurements of eggs and hatchlings of Eutropis carinata (NH: not hatched; SVL: Snout-Vent length; TL: Tail length)

Acknowledgments The authors wish to thank Indraneil Das, Enrique La Marca and Anslem de Silva for reviewing the manuscript and Mohomed M. Bahir for his valuable comments and Mendis Wickramasinghe (SLHS) is also acknowledged for his comments. Finally, we would like to thank Thasun Amarasinghe (TNCS) and Niranjan Karunarathna for their supports. Literature Cited Amarasinghe, A. A. T., 2009. Editorial; An introduction to Taprobanica. Taprobanica, 1 (1): 1. Das, I., A. de Silva, and C. C. Austin, 2008. A new species of Eutropis (Squamata: Scincidae) from Sri Lanka. Zootaxa, 1700: 35–52. Das, I. and A. de Silva, 2005. Photographic Guide to Snakes and other Reptiles of Sri Lanka. New Holland Publishers (UK) Ltd., London: 144. Deraniyagala, P. E. P., 1953. A Colored Atlas of some vertebrates from Ceylon, Vol. 2, Tetrapod Reptilia, National Museums of Sri Lanka, Colombo: 101. de Silva, A., 1996. The Herpetofauna of Sri Lanka: a brief review. Graphic Land, Kandy. Published by Author: 99. de Silva, A., 2006. Current status of the reptiles of Sri Lanka. In: Fauna of Sri Lanka: Status of Taxonomy, Research and Conservation. 134–163. C.N.B. Bambaradeniya (Ed.). The World Conservation Union, Colombo, Sri Lanka and Government of Sri Lanka.

No. Egg size (mm) Hatchling size (mm)

Length Width SVL TL 1 15.2 9.8 28.4 42.3 2 13.7 11.6 26.2 36.5 3 16.3 14.8 29.1 38.7 4 18 9.7 30.2 46.3 5 15.1 11.9 28.1 41.7 6 15.5 11.3 28.3 42 7 16.2 9.9 28.8 43.5 8 15.1 9.8 NH 9 16.5 9.2 29.1 42.4 10 15.7 10.6 28.2 41.1 11 15.5 9.7 NH 12 17.1 9.9 29.8 43.7 13 14.7 9.6 27.4 37.7 14 15.5 9.8 29.1 41.7 15 15.8 9.5 29.7 42.1 16 14.9 9.7 27.2 38.3 17 15.8 9.6 28.6 42.3 18 15.2 9.8 NH 19 15.4 9.7 28.2 41.6 20 17 9.4 29.8 42.4 21 14.3 11.2 27.2 37.6 22 13.8 10.7 NH 23 15.8 11.2 NH 24 15.1 10.5 28.4 42.2 25 15.7 10.2 28.8 41.4

Mean 15.6 10.4 28.5 41.3

65



SOCIAL BEHAVIOURS OF CAPTIVE Trachypithecus cristatus

(MAMMALIA: CERCOPITHECIDAE) IN THE NATIONAL ZOOLOGICAL

GARDENS OF SRI LANKA


A. A. Thasun Amarasinghe 1,2

, W. Madhava S. Botejue 1 and Lee E. Harding

3,4

1 Taprobanica Nature Conservation Society, 146, Kendalanda, Homagama, Sri Lanka

3 SciWrite Environmental Sciences Ltd. 2339 Sumpter Drive, Coquitlam, British Columbia, Canada

Corresponding authors: 2 [email protected];

4 [email protected]

Abstract Trachypithecus cristatus (Silvered Leaf Monkey or Silvered Lutung) occur on the Malay Peninsula, Sumatra, Borneo and nearby islands and live mainly in uni-male, female-bonded groups. This study presents

preliminary data on social behaviours for Silvered Leaf Monkey in captivity. Behavioural observations were

conducted on one group (one adult male, four adult females, three juvenile males and four juvenile females).

Data were collected on 30 of the 60 calendar days at the National Zoological Gardens, Dehiwala, Sri Lanka.

All behaviours including agonistic and dominance-related, locomotion, feeding and infant-related

behaviours were included in the analysis. Unlike in wild populations, we found a clear dominance hierarchy

among females.

Key words: Trachypithecus cristatus, Captive Behaviour, Dominance, Hierarchy, Langur, Sri Lanka.

Introduction Lutung, Langur, Leaf Monkey and Surili are

common names for certain Asian and East Indian

long-tailed, slender monkeys in the genera

Trachypithecus, Semnopithecus and Presbytis.

Lutung and Surili are Malay words that apply to the

Southeast Asian genera Trachypithecus and

Presbytis, respectively, and may be preferable to

Langur, a Hindi word that should perhaps be

restricted to Semnopithecus, a mainly south Asian

genus (Bernstein 1968; Brotoisworo 1979;

Geissmann et al., 2004; Osterholtz et al., 2008;

Weitzel & Groves 1985). All have heavy eyebrows



66

SOCIAL BEHAVIOURS OF CAPTIVE Trachypithecus cristatus IN THE NATIONAL ZOOLOGICAL GARDENS OF SRI LANKA


and, often, beardlike hairs on their chins and body

heavily built (Phillips, 1980). Langurs are

distributed in Sri Lanka, Western Ghats, Bhutan,

Assam and Bangladesh, while Lutungs and Surilis

occur throughout Indo-China, the Malay Peninsula, Archipelago, Bali, and Lombok (Francis, 2008).

Silvered Leaf Monkeys (Trachypithecus cristatus)

occur on the Malay Peninsula, Sumatra, Borneo and

nearby islands. They live mainly in uni-male,

female-bonded groups. Trachypithecus neonates are

orange. Like other leaf monkeys, they are arboreal,

move among the branches by both quadrupedal

locomotion and semi-brachiation (Fleagle, 1977a),

live in troops and feed on vegetation. Arboreal

animals occupying upper story vegetation, which

provide an unobstructed view of the surroundings,

can easily avoid contact. It is therefore interesting

that males regularly sought other males to display

against. Although encounters were frequent, the

cost to the participants was minimal because

physical contact and injury rarely occurred (Poirier,

1968). T. cristatus shows a low level of aggression

and frequent sociosexual, gestural, and vocal

interactions within the social group, relative to other

leaf monkeys (Roonwal & Mohnot, 1977).

Trachypithecus species have relatively short lower

legs and forearms (Washburn 1944), differences in

musculature, and differences in the pelvis (Tague,

1993) compared to Presbytis species. These

adaptations to more quadrupedal locomotion and

less brachiation (Fleagle, 1977a, 1977b) reflect the feeding preferences of Trachypithecines for the

middle and upper canopy, but not as high up or as

far out on slender branches as Presbytis (Curtin,

1977).

Langurs, Lutungs and Surilis are often called Leaf

Monkeys because of their diet, although typically

over one-third of what they eat is seeds, buds,

blossoms, and fruit. Like all colobines, they have a

large, sacculated, ruminant-like stomach with

microbial fermentation, an adaptation to a

folivorous diet (Strasser & Delson 1987). Although

among the smaller of the Trachypithecus and

Presbytis species, T. cristatus had the largest

gastrointestinal tract (Chivers, 2001), a reflection of

its higher intake of leaves and less fruit. Leaves may

comprise up to 91% and fruit only 9% (Hock &

Sasekumar, 1979), the highest and lowest

percentages, respectively, of all colobine species for

which data were available (Caton, 1999; Yeager & Kool, 2000). Most seed eating occurred during the

period of maximum fruit production and fewer

small-seeded, animal-dispersed fruits were eaten at

other times. These included sweet, fleshy fruits,

which most other colobines tend to avoid. Young

leaf intake was greatest in months when fruit intake

was low. Mature leaves were rarely eaten (Glyn,

1991; Kool, 1993). Although some Presbytis species have been seen to consume animal prey

(Goodman, 1989), virtually 100% of the diet of

these monkeys is plants.

T. cristatus has a polygynous, cooperative-breeding

mating system. Single male/multifemale group sizes

of about 10–40, as well as singles and smaller

multi-males groups are the norm; in wild

populations, communal nursing and other parental

care is common and there is no evidence of a female

hierarchy (Bernstein, 1968; Furuya, 1961; Roonwal

& Mohnot, 1977; Wolf & Fleagle, 1977).

Occasionally, a male from an all-male unit or an

individual male will replace the male of a

male/female group. Wolf & Fleagle (1977) reported

that, three months after a male take-over, all of the

dependent infants in the group had disappeared and

were assumed to have been killed by the new

dominant male. This incident of supposed

infanticide by an incoming male has been widely repeated, although actual infanticide was not

observed (as it has in other Trachypithecus species–

Brotoisworo, 1979; Li & Rogers, 2004; Zhao &

Pan, 2006 and Semnopithecus– Hrdy, 1974). It

remains uncertain how often and under what

circumstances infanticide occurs in T. cristatus

(Newton & Dunbar, 2001; Zhao & Pan, 2006). The similarity of these observations to observations of

male-takeovers and subsequent infanticide in Indian

and Ceylonese Langurs (Hrdy, 1974) suggests a

similar pattern of social change among all Langurs,

which live in one-male groups. Borries et al. (1999)

demonstrated the adaptive benefit (i.e., more

progeny) of infanticide by an incoming male

Hanuman Langur (Semnopithecus vetulus) after he

takes over a troop from another male. The

occurrence of male replacement suggests instability

of multi-male organization in bisexual troops.

Moreover, the different mortality rate between

males and females and the unequal sex ratios forced

by the formation of one-male troops, maintained

high-tension levels among males competing for

females (Kunkun, 1986). Genital swellings, which

occur in some Cercopithecinae primate species that

live in multi-female groups and are associated with

oestrus, were examined in adult females by

Shelmidine et al. (2007). They found that vulval swellings occurred minimally, did not increase

during the period of receptivity, and were not

attractive to males.

67



Materials and Methods Study Area: The National Zoological Gardens

(NZG) is approximately 23 acres in extent

(Karunarathna et al., 2008). It is located at a mean

elevation of 20 m above sea level and belongs to the

lowland wet zone of Sri Lanka. The NZG is situated

at the intersection of 6o 50` N and 79

o 54` E,

approximately 2 km away from Dehiwala town and

11 km away from Colombo (Weinman, 1957). The

NZG receives <2000 mm of mean annual rainfall,

with mean annual temperature ranging from

approximately 27.1 oC to 29.4

oC (Somasekaran,

1988). The NZG consists of several habitat types

that can be categorized as man-made small

grasslands, scrublands, several small ponds, home

gardens and large shady trees (i.e. Ficus sp.

Tabubbia sp.).

Study Group: A one-male/ multi-female group of

Silvered Lutungs housed at the National Zoological

Gardens in Sri Lanka was studied. The animals

resided in a 25 m2 naturalistic outdoor enclosure

during the whole day. At the beginning of the study,

the group had one adult male (Mm*), four adult

females (FmL1 & FmL2 = lactating, FmP = pregnant

and Fm*), three juvenile males (Mj

1, Mj

2, Mj

3) and

four juvenile females (Fj1, Fj

2, Fj

3, Fj

4); therefore,

data for twelve individuals were entered in to the analysis. Individual identifications were based on

the shape of the crest, the eye rings and the white

muzzle, as well as the shape of a depigmented skin

area on the belly. Since one female gave birth to an

infant, at the end of the study the group had thirteen

individuals.

Data Collection and Analysis: The group was

observed during half- or all-day sessions during the

normal zoo exhibit times, from 08:00 hr to 18:00 hr

from 01 September through 30 October 2007

(during 30 of the 60 calendar days). The total

recording time was 180 hr. All the behaviours were

assessed via direct observation. Within the group,

observation time was identical per individual.

Additional behaviours were noted whenever they

were observed (ad libitum sampling method). All

the feeding, infant-related, agonistic, locomotor,

contact, self-directed & solitary behaviours and

vocalization were included this analysis. Observations were categorized into wet and dry

periods.

We determined the overall rates of each behavior (N

per individual observation hour). Dominance

hierarchies were constructed based on all focal

agonistic behaviour and ad libitum sampling.

Results Feeding Behaviours: At 10:30 hr and at 14:00 hr,

zoo staff feed the animals. Generally, a mixture of

fruits (papaw, pineapple, mango, plantains, water

melon, lemon, wood apple, apple), sugar cane,

sweet potato, cucumber, long beans, gram (a kind of

pulses), and cabbage are supplied at the morning

meal. In addition, a mixture of rice and bread is

supplied for the afternoon meal. Rarely, green

leaves and Ipomoea aquatica are supplied. Among

these foods their favorites are gram, cabbage, long

beans, green leaves and Ipomoea aquatica. There

was a massive competition among the group to take

above five foods.

Normally, the food tractor comes near to the cage at

10:00 hr and 13:30 hr. As these times approached,

all the animals, especially immatures, were active

and the male remained near the gate and checked

frequently to see whether the food had been put into

the feeding tank (Fig. 1). The other members waited

on the hood of the separated room (Fig. 2). When

we played a sound of a tractor during non-feeding

times, no one reacted to it. When we played it

during feeding time, however, all the animals

reacted immediately to the sound without any

diffidence.

Fig. 01: male checking food

68



When the food tractor reached them, most of

immatures made a noise that we represent as

krēēkk. After the tractor came, these immatures

made a sound that we represent as chrēēss. At this

time most of individuals were active and were moving around the cage. While feeding, the male

took out some foods first and moved aside to eat.

Then lactating females approached the tank,

selected foods, and ate while sitting there. After that

other mature females and older immature males

jumped on and strewed all the food around while

quarreling. They competed for gram, cabbage and

long beans. During this quarreling the immature

females took an opportunity to collect food and then

went away to eat. In addition, they were gluttonous

for green leaves and at that time they frequently

took leaves out of the mouth of other members.

Fig. 02: others waiting for food

The monkeys, and especially the immatures, reacted

quickly whenever visitors offered food, a frequent

occurrence even though prohibited. Whenever visitors came to near the cage, immatures suddenly

raced to the front and begged for food (Fig. 3).

Therefore, most of the immatures as well as adults

appeared addicted to cooked food.

Fig. 03: reaction to the human for food

Agonistic Behaviour & Dominance Relationships

in Adults: During the focal observations, 98 (0.54

interactions per hour) agonistic interactions were

observed; we observed displacement on 22 (0.12

interactions per hour) occasions, and individuals taking food from others on 76 (0.42 interactions per

hour) occasions. 78% of these agonistic interactions

(displacement and taking food from others)

occurred during feeding period. During the

observations, Mm* was barking on 6 (0.03

interactions per hour) vocalization interactions.

Based on these relationships (who consistently did

what to whom), we constructed a dominance

hierarchy among females and immature males. The

youngest mature female (Fm*) already acquired a

medium rank, while the other four of the immature

females (Fj1, Fj

2, Fj

3, Fj

4), were still at the bottom

of the hierarchy. The older immature male (Mj1)

was above the female dominance hierarchy, but

below the adult male (Mm*). The two younger

males (Mj2, Mj

3) were still low ranking, but above

the same-aged immature females. In addition, some

agonistic behaviours of the dominant male against

visitors were recorded (Fig. 4).

Fig. 04: an agonistic behaviour of the male against

visitors

Infant Related Behaviour: During the observations,

Fj4 nipple-contacted 154 times (0.85 interactions per

hour). FmL2

withdrew nipple contact 32 times (0.17

interactions per hour), Mj3

nipple-contacted (Fig. 5)

114 times (0.63 interactions per hour) & FmL1

withdrew nipple contact 40 times (0.22 interactions

per hour). We observed FmL2 attempt to take her

infant Fj4 away from others on 40 (0.22 interactions

per hour) occasions; out of these 40 occasions, 12

were rescues. FmL1

attempted to take her infant Mj3

away from others on 16 (0.08 interactions per hour) occasions; out of these 16 occasions, 8 were

rescues.

69



Most lactating females trended to feed their infants

during 08:00 – 09:00 hr (before the morning feeding

time), 11:00 – 12:00hr (after the morning feeding

time) and 13:00 – 14:00 hr (before the afternoon

feeding time). In addition, females spent a total of 32 hours (18% of the total observation time) feeding

their infants.

Fig. 05: nipple contact behaviours

The younger immature Fj4

tried to play with other

immatures (Fig. 6) and also tried to imitate them.

However, most of time the mother FmL2

repressed the infant and took it away. During this time the

infant made the chrēēss vocalization. The infant

also made this sound while it had been in a

dangerous situation. On these occasions, FmL2

reached quickly for the infant and took it away from

the danger.

Fig. 06: hanging and climbing behaviours

Locomotor Behaviour: During the focal

observations, (1 hr for each juvenile) we recorded

1116 locomotory behaviours for 7 individuals

(individually ~29 actions per hour), in wet weather

conditions and 3176 for 7 individuals (individually ~ 82.4 actions per hour) in dry weather conditions.

Here we considered only locomotor behaviours of

juveniles for this analysis and omitted all other

behaviours (feeding, agonistic, infant-related,

contact behaviour, self-directed & solitary and

vocalization).

During both wet and dry days, the group was active

from 10:00 hr to 11:30 hr and 14:00 hr to 15:00 hr.

Their feeding time also overlapped at 10:30 hr and

14:30 hr. Before 08:00 hr, between 13:00 – 13:30 hr

and after 17:30 hr the immatures as well as adults

were significantly inactive. During the observations,

adults never showed suspensory locomotions (i.e.,

brachiation) or hanging behaviours, while juveniles

mostly showed suspensory locomotions.

In addition to the solitary behaviours (Fig. 7), a few

grooming behaviours (Fig. 8) and scratching

behaviours were recorded. These observations were not entered in to the analysis due to the lack of

frequency.

Fig. 07: solitary behaviours

70



Fig. 08: grooming behaviours

During the observation period, the male and FmP

sunbathed for about 10 minutes, evidently to heat

their bodies, while clinging to a granite rock (Fig.

9). They rarely showed this behaviour after the

afternoon meal.

Fig. 09: sun bathing behaviours

Discussion and Recommendations The diet of this captive population differed

markedly from that of wild populations, in which

much higher proportions were leaves, including

mature leaves (Roonwal & Mohnot, 1977; Hock &

Sasekumar, 1979). They spent approximately 20%

of feeding time foraging on Moraceae species. In a

field study on the closely related Trachypithecus

auratus, teak (Tectona grandis) was the top food

species (Kool, 1993), but in our study, the captive

leaf monkeys never got T. grandis or other native

foods. However they liked to eat rarely given green

leaves, Ipomoea aquatica, gram, cabbage and long

beans. Therefore we assume the taste of given

green-leaves; l. aquatica, gram, cabbage and long

beans may be similar to those that are eaten in

natural habitats. These more natural foods may also

be healthier, considering the physiological adaptations of T. cristatus to a highly folivorous

diet; and there was no obvious evidence of

malnutrition or vitamin deficiency. The monkeys’

reactions to the tractor sound at feeding times, but

not at other times, suggests that this may be a

common, learned response in captive populations.

Unlike in wild populations (Bernstein, 1968; Furuya

1961; Medway, 1970; Roonwal & Mohnot, 1977;

Wolf & Fleagle, 1977), there was a clear dominance

hierarchy among the females. The dominance

hierarchy of the group, in order from the bottom,

was Fj1, Fj

2, Fj

3, Fj

4 < Mj

2, Mj

3 < Fm

P < Fm

L1, Fm

L2

< Fm* < Mj

1 < Mm*. Here the Fm

L1, Fm

L2 was at the

medium rank, just below the top male during the

feeding time. During feeding times, we could

identify the hierarchical order from the bottom; Fj1,

Fj2, Fj

3, Fj

4 < Mj

1, Mj

2, Mj

3 < Fm

*, Fm

P < Fm

L1,

FmL2 < Mm*. Therefore we suppose that the group

(especially Fm* and Mj

1) allowed Fm

L1, Fm

L2 to

acquire foods first because they were feeding

infants. The presence of a female hierarchy in this

study suggests caution in extrapolating data from

captive to wild populations. Alternatively, a female

hierarchy might have been overlooked in wild

populations and might be detected with more

careful observation of marked or otherwise identifiable individuals.

According to agonistic behaviours and dominance

relationships, the immature male (Mj1) was at the

very top of the female dominance hierarchy, but

below the adult male [alpha (α) male] (Mm*).

Therefore we imagine that the immature male (Mj1)

would become the beta (β) male in the near future.

Although males were dominant above same-aged

females in this study, it is possible that the

dominance was based on size, rather than gender,

since T. cristatus is moderately sexually dimorphic

(c.f. Hemelrijk et al., 2008).

Most lactating females trended to feed their infants

during before the morning feeding time, after the

morning feeding time and before the afternoon

feeding time. Lactating females refrained from

feeding their infants before having their own meals.

Therefore we believe there is a synchrony between

the feeding time and lactating time.

During focal observation of the adult male, we

could observe some grooming and unsuccessful

copulation behaviours with mature female (Fm*).

According to Shelmidine et al. (2007), although genital swellings were most common during the

breeding period, there was no strong correlation

with oestrus and no particular attraction to the

71



males. However, the vulva of Fm* was not swelled

in this period, suggesting that she was still not

sufficiently mature for a successful copulation.

Finally we would like to recommend giving favorite and more natural foods to this species, not only to

minimize refusing & wasting, but to ensure a

healthy diet. In addition, further research should be

continued for longer duration with more detail on

silvered lutungs. As well, other behavioural studies

of animals at the national zoo should be promoted

as they may help with ex-situ conservation. Most

importantly, visitors should be strictly prohibited

from feeding animals and giving non-digestible

materials that cause diseases and harm. This

restriction should be combined with education to

encourage visitors to love animals and observe their

behaviours without harm.

Acknowledgements The authors wish to thank Professor Colin Groves

for reviewing the manuscript and Carola Borries

and Mohomed M. Bahir are also acknowledged for

providing valuable comments on the earlier draft.

We also thank Damith Wijewardena (TNCS),

Gayan Wijethunge (TNCS), Dinesh Gabadage

(TNCS) and Sanjeewa Lakmal who helped in

diverse ways to enrich this work.

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