Reptiles of the Solomon Islands (Faunistica)

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uring the 26 years Michael McCoy lived in the Solomons, he visited virtually every island in the archipelago and is widely regarded as one of the foremost authorities on the reptile fauna of the Solomons. He has published several books, worked as a consultant on a number of television documentaries, and his photographs have appeared widely in regional and international publications. He now lives in Australia.

Pensoft Series Faunistica No 57 ISSN 1312-0174

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REPTILES OF THE SOLOMON ISLANDS Michael McCoy

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his book provides comprehensive accounts of 86 reptiles, which represent all the currently known species occurring in this island archipelago. Most importantly, the natural histories of the living animals are given in detail, and almost all species – including many colour variants – are depicted in large, colour photographs. Designed as a field guide, the book is compiled in such a way as to be of use not only for professional biologists working in the field or museum, but also for the interest of amateur naturalists and travellers in Solomon Islands. In fact, anyone with an appreciation for the natural history of this group of beautiful South Pacific islands will find this book a worthwhile addition to their library.

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REPTILES OF THE

SOLOMON ISLANDS

Michael McCoy

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REPTILES OF THE SOLOMON ISLANDS

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REPTILES OF THE

SOLOMON ISLANDS WRITTEN AND PHOTOGRAPHED BY

Michael McCoy

Sofia–Moscow 2006

REPTILES OF THE SOLOMON ISLANDS Written and photographed by Michael McCoy

First published 2006 ISBN-10: 954-642-275-4 ISBN-13: 978-954-642-275-0 Pensoft Series Faunistica No 57 ISSN 1312-0174

© PENSOFT Publishers All rights reserved. No part of this publication may be reproduced, stored in a retrieval system or transmitted in any form by any means, electronic, mechanical, photocopying, recording or otherwise, without the prior written permission of the copyright owner.

Pensoft Publishers Geo Milev Str. 13a, Sofia 1111, Bulgaria [email protected] www.pensoft.net

Printed in Bulgaria, September 2006

CONTENTS 7

Foreword

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Acknowledgements

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Map of Solomon Islands

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Introduction

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Crocodiles, Family Crocodylidae

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Marine Turtles, Family Cheloniidae

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Leatherback Turtle, Family Dermochelyidae

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Agamid Lizards, Family Agamidae

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Geckos, Family Gekkonidae

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Skinks, Family Scincidae

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Monitor Lizards, Family Varanidae

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Blind Snakes, Family Typhlopidae

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Boas, Family Boidae

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File Snakes, Family Acrochordidae

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Colubrid Snakes, Family Colubridae

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Elapid Snakes, Family Elapidae

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Literature Cited and Further References

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Glossary

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Island Check Lists

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Index of scientific names

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FOREWORD Before Wau Ecology Institute published the first edition of Mike McCoy’s field guide to the reptiles of Solomon Islands in 1980, our herpetological knowledge of this remote corner of the tropical South Pacific was very scant, with the few available scientific papers – most decades old – widely scattered in diverse journals. Arriving in the Solomons in 1969, McCoy began field studies and photography of the reptile fauna. In the mid 70s he spent several months working through the preserved collection of Solomon reptiles at the Australian Museum. These field and museum studies later provided the basis for McCoy’s 1980 book. During the 26 years Mike McCoy lived in the Solomons, he visited nearly all the islands in the archipelago and is widely regarded as one of the foremost authorities on the reptile fauna of the Solomons. He is also an extraordinarily gifted photographer with a keen eye for detail, an indefatigable field worker and an old and close friend. Since the publication of the first field guide in 1980, he has continued his studies, making many additional discoveries, and this fine book is the culmination of those efforts. The Solomons’ extraordinary reptile fauna includes a wealth of small geckos and skinks, and also includes such oddities as Tribolonotus, represented in the Solomon Islands by several species which have highly carinate or spinose scales, produce single young and practice parental care. And there’s Corucia zebrata, the world’s largest skink; highly arboreal and prehensile-tailed, it is a Solomons endemic and gives birth to a young nearly a third the size of the adult. There are endemic venomous elapid snakes, innocuous, fossorial blind snakes, several species of boas and the ubiquitous saltwater crocodile. For the herpetologist, the Solomons are a unique living laboratory. By studying such groups as, for example, the wide-ranging skink genus Emoia FOREWORD

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throughout the islands, we have been able to gain valuable insights into the mechanisms of reptilian phylogeny; how insular reproductive isolation gives rise to the evolution of new species, and the influence of geological history on evolutionary diversification. All but a few of the reptile genera found in the Solomon Islands are shared with neighbouring New Guinea, but there has been an extraordinary radiation of species in association with the complex geological history of the southwest Pacific region. More than 80 species of reptiles are now known from the Solomon Islands and future work may well increase this number to over a hundred. ALLEN ALLISON Ph.D. Vice President, Science Bishop Museum, Honolulu

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FOREWORD

ACKNOWLEDGEMENTS I would like to thank the many Solomon Islanders who have collected reptiles for me and with me during the 26 years I lived in the Solomons. Especially I thank Peter Beata, Aidan Bu’uga, Gabriel Feroa and Davy Maerufa. For scientific information regarding Solomons reptiles, and providing me with published literature, I thank Allen Allison, the late Walter Brown, Harold Cogger, Allen Greer, Glenn Shea and George Zug. I would also like to thank Harold Cogger for allowing me ready access to the preserved reptile collection at the Australian Museum during his tenure as deputy director. Allen Allison and George Zug read the draft manuscript of this work and offered helpful suggestions and criticisms.

ACKNOWLEDGEMENTS

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Map of Solomon Islands. The inset map of Santa Cruz and the Eastern Outer Islands is at the same scale as the main map.

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MAP OF SOLOMON ISLANDS

INTRODUCTION The Solomon Islands archipelago is situated in the tropical South Pacific to the east of New Guinea and the northeast of Australia. The group is made up of the major islands of Bougainville, Choiseul, the New Georgia Group, Isabel, Guadalcanal, Malaita and Makira and numerous smaller islands and atolls. The terms Solomon Islands, or Solomons, as used within the context of this work refer to the island archipelago and also include the Santa Cruz Group and the Eastern Outer Islands (about 500 km east of the main archipelago) that comprise Temotu Province of the nation of Solomon Islands. The island of Bougainville, while politically part of Papua New Guinea, is geographically and zoogeographically a major component of the Solomons archipelago. These islands lie between 50 and 120S and 1540 and 1700E and have a total land area of around 37,000 kms2. The larger islands of the Solomons are basically alike in having a central mountainous spine (usually of 1000 m elevation or more) falling away through a series of foothills to the coast. Rainfall is high throughout the group with the annual average around 3000–4000 mm. Rivers and creeks are numerous on all the major islands and on many of the smaller ones. Tropical rainforest is the predominant vegetation, though both large and small islands are fringed with extensive mangrove growth in many areas. Bougainville, the largest island of the group, has active volcanoes. Guadalcanal is unique with its extensive grass plains on its northern coast. Human habitation is largely confined to the coasts. In rural areas extensive areas of forest adjacent to villages are cleared to accommodate the traditional food gardens. Distances between islands vary, Rennell is separated from Guadalcanal by more than 200km of open sea, while Maramasike (Small Malaita) is separated from Malaita by as little as 300–400 m. Generally though, the main islands are separated by an average of around 50–100km. The climate is tropical with temperatures normally within the range of 220–340C on an annual basis. From May to October the southeast tradewinds blow, gusting at times up to 30 knots INTRODUCTION

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or more. During this dry season the days are mostly fine with the occasional rain squall. From November to April the wind is from the northwest. This is the wet season or monsoon and is characterised by frequent heavy rain and varying wind intensity. Cyclones, with destructive winds and waves, frequently develop during the later months of the monsoon. The Solomon Islands are comparatively recent, dating from mid-Miocene times (about 15 million years ago). Zoogeographically they are towards the end of a migratory path leading down from southeast Asia, through New Guinea and eastwards to the Bismarcks-Solomons archipelago. They are home to an abundant vertebrate fauna and are herpetologically rich with more than 80 species of reptiles, around a third of which are endemic. Although the following work represents all the reptile species known to occur in the Solomon Islands at the time of publication, this account should not be regarded as complete by any means. The interior and mountainous areas of the larger islands have been poorly collected and new species may still be found. Furthermore, largely due to recent DNA studies, it has been established that some animals currently recognised as single species are composite, encompassing several as yet undescribed species. This is certainly true for some skinks of the genus Sphenomorphus, and probably also for some members of genera such as Emoia and Lepidodactylus. This book is essentially a field guide, and is compiled in such a way as to be of use not only for professional biologists, but also for the interest of amateur naturalists and travellers in Solomon Islands, and in fact anyone with an appreciation for the natural history of this group of beautiful South Pacific islands. To identify a reptile using this book, the photographs should be checked initially for provisional identification, followed by reference to the species account for confirmation. In this way a positive identification can usually be made. If certain identification is still in doubt, then the keys should be used. For some reptile specimens, particularly very small animals, a powerful hand lens or microscope may be necessary to determine characters such as scale counts. However, identifying a blind snake (Typhlopidae) on the basis of scale counts and external characters is virtually impossible without a good quality microscope, and the typhlopid keys are intended for laboratory and field use where a microscope is available.

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INTRODUCTION

Each species account is divided as follows: 1. The scientific or Latin name (a binomial of genus and species) and the person or persons who originally described the species (the author or authors). Where the author’s name appears in parentheses it indicates that subsequent revision has resulted in the species being assigned to a different genus from which it was originally described. 2. The distribution. The ranges outside the Solomons are given to indicate the extent of a species’ distribution only and should not be regarded as exhaustive. Within the Solomons, the islands listed represent those localities where that species has been collected and noted accordingly in the scientific literature, or where I have personally found that species. Again, these locality data should not be regarded as complete by any means, as future field studies in the Solomons will inevitably result in new distributional records for many reptiles. Checklists of terrestrial species for all the major islands and many of the smaller islands are provided at the back of the book. 3. The description of body form, scalation and scale counts will in most cases allow for positive identification of live or preserved specimens, or for the few animals that are not illustrated. The glossary at the end of the book can be consulted by those not familiar with the terms used. The average size given for each species does not represent a statistical mean, it is an approximation of the average adult size. The scale and lamellae counts represent only the recorded minimum and maximum counts to date and should not be regarded as absolutes. These descriptions are derived from the available scientific literature and also from Solomons reptile specimens that I have collected and examined myself or are stored in the preserved collections of various natural history museums. 4. The colour and pattern are given as aid to identification, as is some idea of the variability of these features. Such variability can be seen in the photographs of a number of species. Because the colour and pattern are given in broad terms only, a certain flexibility must be allowed if an individual does not strictly conform to the colour and/or pattern in the species account or the illustrations. Variation in colour and/or pattern often occurs in the same species from different islands, and in some cases in populations from the same island, and such variation may be considerable. Colour descriptions are of live animals unless otherwise indicated. INTRODUCTION

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5. Natural history notes. In this section a brief account is given (as far as is known) of the animal’s habits and lifestyles. Almost all of these accounts are based on my own observations of free-living reptiles, and to a lesser extent on captive specimens. In some cases I have included additional information I have received from Solomon Islanders living in rural areas and whose observational expertise I respect. In the first print edition of this book, published in 1980, I wrote in the Introduction: “The conservation of the environment should therefore be the concern of any far-sighted and responsible government. Exploitation of the environment for short-term economic gains is not only morally wrong, it also deprives future generations of the aesthetic benefits of an undisturbed ecosystem”. That the Solomons’ natural environment with its myriad species of plants and animals would flourish under the guardianship of responsible and caring governments has sadly proved to be nothing more than naïve wishful thinking. In the more than 25 years since I wrote the above words, much of the lowland areas of Solomon Islands have been subject to severe environmental degradation. A burgeoning human population has rapidly and extensively expanded subsistence agriculture into previously untouched rainforest, and coupled with the virtually unchecked logging activities by totally unprincipled and unscrupulous foreign operators, has reduced much of the pristine forests of the islands to ecological disaster areas. Additionally, the depredation on select reptile species - notably Corucia zebrata, Varanus indicus and the Candoia species - by collectors supplying the U.S. and European pet trade with literally thousands of animals annually have left the long-term survival of wild populations of many Solomons reptiles in doubt. Although this insidious trade in wildlife was banned by the Solomon Islands government in 2005, the ban is poorly enforced and the illegal export of reptiles, and also amphibians and birds, continues.

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INTRODUCTION

CROCODILES, Family CROCODYLIDAE Crocodiles and their relatives are an ancient group of reptiles. They occur throughout tropical regions around the world. Crocodilians differ from other reptiles in having a four-chambered heart (the ventricle is completely divided). The external ear and nostrils are valvular, and together with their powerful tail, give ready testimony to their aquatic existence. Genus Crocodylus Laurenti, 1768 Snout relatively long and broad. The fourth mandibular tooth fits into a notch on the upper jaw and is more or less visible when the mouth is closed. SALTWATER OR ESTUARINE CROCODILE Crocodylus porosus Schneider, 1801 Fig. 1 Distribution: From India through southeast Asia to New Guinea and its island archipelagos, northern Australia, Solomon Islands, Vanuatu. Very occasional individual animals, not representatives of resident populations, have been recorded from Palau in the Caroline Islands, Nauru and Fiji. In the Solomons it occurs on all the larger islands and many of the smaller islands of the group. Description: Average total length around 2.5m although it reputedly grows to 6m. Snout fairly broad, about 2x as long as wide, 17 to 19 teeth on each side of the upper jaw. Usually 4 enlarged keeled nuchal scales, separated from the smooth occipital region by several rows of smaller granular scales. Colour: Grey or grey-brown above, often with irregular lighter and darker markings. Ventrally creamy white. Natural History Notes: The species inhabits mangrove areas and tidal estuaries, also creeks and rivers, swamps and on occasion, the open sea. During the day crocodiles occasionally come ashore to bask, though they will immediately take to the water if they are disturbed. At night larger crocodiles CROCODILES, Family CROCODYLIDAE

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often move around on land, up to 100m or more away from the water. They feed on a variety of vertebrates and invertebrates such as crabs, fish (including sharks), frogs, birds, other reptiles including smaller crocodiles, mammals such as rats, flying foxes and very occasionally pigs and calves. There have been human fatalities attributed to this crocodile in the Solomons. I have had some opportunity to observe wild crocodiles on the island of Aliiti in the Olu Malau group off Makira in the eastern Solomons. On Aliiti a large colony of flying foxes Pteropus sp. live in the trees surrounding the small lake inhabited by the crocodiles. During heavy rain the bats move low in the trees and the crocodiles wait in the water below and grab any bat that comes within reach. On Malaupaina Island in the Olu Malau group I have seen evidence of crocodiles digging in the sand above the high water line, presumably for crabs. When nesting, a large nest of leaves, sticks and grass is constructed by the female crocodile and the eggs are laid in this and guarded by the female until they hatch. No detailed information is available for nesting times or egg numbers in the Solomons; it is not known if crocodiles have a well defined breeding/nesting season in these islands. A 1988 survey estimated the total crocodile population of the Solomons to be less than 1,000 animals with the largest breeding population (about 700 animals) occurring in Lauri Lagoon on the southern coast of Guadalcanal. In 2005 anecdotal information I received from local people on Guadalcanal, Ngela and Malaita indicated that crocodile populations on those islands were increasing. Some popular travel guidebooks on the Solomons mention a second “harmless” species of freshwater crocodile occurring in rivers. This is an incorrect and potentially dangerous piece of misinformation. There is only one species of crocodile in the Solomons, the saltwater crocodile, and even small specimens are far from harmless.

MARINE TURTLES, Family CHELONIIDAE Marine turtles occur throughout the tropical and subtropical seas of the world. All species are oviparous, the female coming ashore to lay her eggs in a hole scooped out in the sand well above the high water line. Their forelimbs are paddle-shaped and one or 2 claws are present on the forelimbs. The carapace is well developed and is comprised of a series of large horny shields. The neck is short and the ability to retract the head into the shell is only partial. They feed on a wide variety of marine organisms, both animal and plant.

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MARINE TURTLES, Family CHELONIIDAE

Vertebrals

Costals

Marginals

Plates of the carapace of a cheloniid turtle (Lepidochelys olivacea)

KEY TO THE SPECIES OF SOLOMON ISLANDS TURTLES 1. Carapace covered with horny shields, forelimbs with claws: 2 – Forelimbs without claws, carapace covered with leathery skin with 7 longitudinal ridges: Dermochelys coriacea 2. Four costal plates: 3 – Five or more costal plates: 4 3. Two pairs of prefrontal scales on the head, upper jaw projecting and beak-like: Eretmochelys imbricata – A single pair of prefrontals, upper jaw not hooked or beak-like: Chelonia mydas 4. Head very large, rarely more than 5 costal plates, usually 3 inframarginals on the bridge without pores: Caretta caretta – Head moderate to large, usually 6 or more costal plates, 4 inframarginals on the bridge usually with pores: Lepidochelys olivacea

MARINE TURTLES, Family CHELONIIDAE

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Genus Caretta Rafinesque, 1814 A monotypic genus. Head very large with symmetrical shields. Two pairs of prefrontals. Five costal plates. LOGGERHEAD TURTLE Caretta caretta (Linnaeus, 1758) The Indian Ocean and Pacific Ocean form is generally referred to the subspecies gigas. Distribution: Throughout the tropical and subtropical Indian, Pacific and Atlantic Oceans. Description: Average carapace length around 1m, reputedly growing to nearly 2m. Head very large, particularly so in adult specimens. Two pairs of prefrontals. Upper jaw not hooked. Carapace somewhat elongated and heart-shaped. Five costal plates (rarely 6), marginals usually 12 to 13 but may vary from 11 to 15, most commonly 3 inframarginals (without pores) on the bridge. Colour: Carapace reddish brown, often with darker variegated irregular markings. Head usually reddish brown above, paling on the sides. Limbs and tail dark brown above, fading to yellowish cream towards their edges and below. Plastron cream to yellowish. Natural History Notes: A carnivorous species rarely seen in the Solomons; only the occasional individual has been reported although most Solomon Islanders living in coastal villages are familiar with this species. Adult Caretta normally feed in depths of less than 100m, although they have been seen feeding in very deep water; a specimen was reportedly observed off Baja feeding on galatheid crabs at a depth of more than 300m (Walter Starck, pers. comm). This species is not known to nest in the Solomons.

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MARINE TURTLES, Family CHELONIIDAE

Genus Chelonia Latreille, 1802 Head with symmetrical shields. A single pair of prefrontals. Four pairs of costal plates. GREEN TURTLE Chelonia mydas (Linnaeus, 1758) Fig. 2 The western Pacific form is generally referred to the subspecies japonica. Distribution: Throughout the tropical and subtropical Indian, Pacific and Atlantic Oceans. Description: Average carapace length 1m. Head moderate. Upper jaw not hooked. A single pair of prefrontals. Carapace broad and heart-shaped. Four costal plates, 5 vertebral plates, 4 inframarginals (without pores) on the bridge. Forelimbs with enlarged irregular scales distally. Limbs with a single claw. Colour: Carapace light brown to greenish brown with radiating or wavy pattern of lighter and darker markings on each dermal plate. The head and limbs are light brown above, cream to white below. The plastron is cream to almost white. Natural History Notes: This turtle is relatively abundant in the Solomons and nests on most islands. The nesting season does not appear to be well defined and nesting occurs throughout the year. In the eastern Solomons I have noticed that there is perhaps more nesting activity in the period November to February. The female digs her nest ashore at night in sandy areas, usually amongst shore vegetation well above the high water line. About 100 eggs are laid in a clutch, the female often coming ashore to the same beach to lay successive clutches over a period of several weeks. The young turtles hatch in about 8 to 10 weeks, always at night. “Explosive” emergence has been well documented in young turtles where the hatchlings emerge from the nest almost simultaneously. Hatchling C. mydas are largely carnivorous though the adults are herbivorous, feeding on sea grasses and algae. At night each turtle often returns to the same place to sleep, usually under coral overhangs at a depth of around 2 to 15m. While diving at night on a reef about 1km offshore from Honiara on Guadalcanal, I noticed on several occasions over a 3 year period, two large turtles, a green and a hawksbill, sleeping together in a large coral cave at a depth of about 12m. It can be reasonably assumed that these MARINE TURTLES, Family CHELONIIDAE

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were the same turtles on each occasion. Their average foraging depth during the day more or less corresponds to the sleeping depth range though I have seen these turtles at a depth of 30 m.

Genus Eretmochelys Fitzinger, 1843 A monotypic genus. Head with symmetrical shields. Two pairs of prefrontals. Four pairs of costal plates. Upper jaw with a distinctly hooked beak. HAWKSBILL TURTLE Eretmochelys imbricata (Linnaeus, 1766) Fig. 3 The Indian Ocean and Pacific Ocean forms are generally referred to the subspecies bissa. Distribution: Throughout the tropical and subtropical Indian, Pacific and Atlantic Oceans. Description: Average carapace length about 700mm though it grows to a little over 1m. Head small to moderate. Upper jaw projecting and forming a distinct hooked beak. Two pairs of prefrontals. Carapace heart-shaped, often with a distinct vertebral keel. The edge of the carapace is serrated posteriorly and the dermal plates overlap except in hatchlings and old adults. Four pairs of costal plates, 5 vertebrals and 4 inframarginals (without pores) on the bridge. Limbs with 2 claws. Colour: Carapace dark brown with mottled lighter and darker patterning. The head is dark brown to black with distinct whitish sutures between the scales. The plastron is dark brown to black in hatchlings and cream to light brown, often with darker markings, in juveniles and adults. Natural History Notes: Similar to Chelonia mydas excepting that it is carnivorous, feeding on a variety of marine animals such as fish, crabs, various molluscs, certain species of sponges and jellyfish, including the highly venomous box jellyfishes. Most feeding activity takes place during the early morning or late afternoon. During the main part of the day the turtle appears more to browse and rest, either on the reef or sometimes at the surface. This species is quite common in the Solomons and nests on most islands. Nesting habits are similar to green turtles. Also like greens, individual hawksbills will return to the same place to sleep each night.

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MARINE TURTLES, Family CHELONIIDAE

Genus Lepidochelys Fitzinger, 1843 Head shields symmetrical. Two pairs of prefrontals. Six or more pairs of costals. Usually 4 enlarged inframarginals on the bridge, each with a single pore. PACIFIC OR OLIVE RIDLEY Lepidochelys olivacea (Eschscholtz, 1829) Fig. 4 Distribution: Throughout the tropical Indian and Pacific Oceans and southern tropical Atlantic. Description: Average carapace length 750mm. Head large. The upper jaw is without a distinct hooked beak. Two pairs of prefrontals. Carapace wide and heart-shaped, flattened dorsally. Six or more pairs of costals (there may be more costals on one side than the other), 4 inframarginals on the bridge, each with a single pore at its posterior edge, 12 to 14 marginals. Limbs with a single claw. Colour: Carapace olive or grey, usually without any distinctive markings. Plastron greenish white to cream. Head and limbs olive to grey above, paling on the sides to creamy white below. Natural History Notes: This species is quite rare in the Solomons. The animal illustrated in fig. 4 is a subadult (about 300mm carapace length) from Wainone Bay on northern Makira. I obtained it from villagers in that area in 1977. They claimed it had been raised from an egg taken from a nest on a black sand beach in Wainone Bay the previous year. If this is correct it is the first known nesting site for this species in the Solomons. The Wainone Bay villagers were unable to recall the month of the nesting but did report that the nest contained more than 30 eggs. A mating pair of this species were found in the sea off Guadalcanal in February 1976. Lepidochelys olivacea is carnivorous, feeding mainly on crustaceans and molluscs.

LEATHERBACK TURTLE, Family DERMOCHELYIDAE This family contains a single species, a highly specialized pelagic turtle. It occurs worldwide in tropical and temperate waters. The carapace lacks epidermal shields; it is made up of a series of small dermal bones embedded LEATHERBACK TURTLE, Family DERMOCHELYIDAE

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in the skin. The limbs, particularly the forelimbs, are very large and lack claws. The head shields are asymmetrical.

Genus Dermochelys Blainville, 1816 See family for description. LEATHERBACK OR LEATHERY TURTLE Dermochelys coriacea (Linnaeus, 1766) Distribution: Worldwide in both tropical and temperate waters. Description: Average carapace length around 1.75m though it grows to over 2m. Carapace covered with leathery skin, elongated and made up, together with the plastron, of a layer of small dermal bones embedded in the skin. On the carapace a number of these bones are enlarged and arranged into a series of 7 longitudinal ridges. The plastron has 5 similar ridges. The forelimbs are particularly large and lack claws. Colour: Dark brown to black above with a scattering of whitish spots and blotches over the head, carapace and limbs. The undersurfaces are yellowish white. Natural History Notes: A pelagic species though occasionally entering bays and estuaries. Leatherbacks feed almost exclusively on jellyfish and have the ability to maintain a body temperature greater than ambient. This turtle is rarely encountered in Solomons waters, though most Solomon Islanders know the species. It is known to nest on several islands of the group including Vella lavella, Choiseul, Tetepare, Isabel, Guadalcanal, Malaita, Ulawa and Makira. Nestings on these islands have invariably occurred on black sand beaches. The dates of nestings are irregular and somewhat infrequent though essentially year round. Dermochelys and its eggs are protected in the Solomons, though female turtles coming ashore to lay in the vicinity of villages are most likely to be killed and eaten by the local people.

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LEATHERBACK TURTLE, Family DERMOCHELYIDAE

LIZARDS, Order SQUAMATA, Suborder SAURIA KEY TO THE FAMILIES OF SOLOMON ISLANDS LIZARDS 1. Nuchal and dorsal crests present, tail more than 300% of SVL: Agamidae – Nuchal and dorsal crests absent, tail less than 300% of SVL: 2 2. Eye covered with an immovable transparent shield, pupil vertically elliptic, dorsal scales very small, granular and juxtaposed: Gekkonidae – Eye with a movable eyelid or if immovable then body scales smooth and imbricate, pupil round: 3 3. Head scales symmetrical, body scales imbricate, at least ventrally if not dorsally, tail more or less round in cross-section: Scincidae – All body scales juxtaposed, tail laterally compressed for most of its length: Varanidae

AGAMIDS, Family AGAMIDAE A diverse group of small to moderate sized lizards commonly called agamids or dragons. They occur in Africa, parts of southern Europe, much of south and southeast Asia, Australia, New Guinea and several islands of the western Pacific. They are characterized by their asymmetrical head scalation and juxtaposed body scales which are often enlarged and spinose. The tongue is broad and fleshy. Nearly all species are oviparous and many species have femoral and precloacal pores. All are essentially active diurnal lizards and occupy a diversity of habitats from arid deserts to tropical forests and some are partly aquatic. Depending on size, they feed on a variety of vertebrates and invertebrates and many species eat some plant material such as blossoms and fruit. Genus Hypsilurus Peters, 1867 Mostly arboreal. Head large and angular with a distinct tympanum. A transverse gular fold is present. Dorsal and nuchal crests are present. Body strongly compressed. No femoral or precloacal pores. Earlier literature AGAMIDS, Family AGAMIDAE

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invariably refers to a single species of Solomons agamid, Hypsilurus (previously Gonocephalus) godeffroyi. Manthey and Denzer (2006) now recognise two Solomon species of Hypsilurus and believe that “true” H. godeffroyi does not occur in the Solomons. KEY TO THE SPECIES OF SOLOMON ISLANDS Hypsilurus – Large size (SVL of adults usually greater than 150mm), a V-shaped mark on the nape of the neck in females and young males, occurring on Bougainville (and probably Shortland Islands and Isabel): longii – Moderate size (SVL of adults usually less than 130mm), no V-shaped mark on the neck in females or males, occurring in the eastern Solomons (Makira, Ugi, Olu Malau and Santa Ana): macrolepis Hypsilurus longii (Macleay, 1877) Distribution: Bismarck Archipelago and possibly mainland New Guinea, Solomon Islands, northern Cape York in Australia. In the Solomons: Bougainville. Populations of Hypsilurus in the Shortlands Group are most likely to be referrable to this species. An indigenous name for Hypsilurus exists on northern Choiseul which is adjacent to Bougainville, but villagers there have told me that these lizards do not now occur in that area. Manthey and Denzer (2006) distinguish H. longii from H. macrolepis partly on the basis of SVL (150mm or greater in H. longii, 130mm or less in H. macrolepis). While collecting reptiles on Isabel Island for the Australian Museum in 1987, Pavel German (pers. comm.) saw, but did not collect, a large Hypsilurus with an estimated SVL of around 250mm. Given its size, this animal was almost certainly H. longii. Three specimens of Hypsilurus from Isabel in the Australian Museum have SVLs ranging from 185mm to 223mm (Ross Sadlier, pers. comm.) and are therefore also referrable to H. longii on the basis of Manthey and Denzer’s definition. Type locality: North Queensland, Australia. Description (adapted from Manthey and Denzer, 2006): SVL of males up to 235mm, females up to 210mm, both usually more than 150mm, tail more than 300% of SVL. A large, robust species. Several scales below the tympanum sometimes strongly enlarged, anterior edge of the gular pouch without enlarged scales. Head scales small, homogeneous, strongly keeled. Temporal scales slightly enlarged, several scattered enlarged scales below the tympanum

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AGAMIDS, Family AGAMIDAE

often forming 1–2 rows. 10 supralabials, 9 infralabials. Gular pouch moderately large, anterior edge without enlarged scales, a weakly developed saw-toothed edge sometimes visible. Continuous vertebral crest running to the base of the tail with a significant indentation on the nape. Base of the tail laterally strongly compressed and a little raised (in old males clearly raised). Colour (in alcohol) (from Manthey and Denzer, 2006): Back and sides with 4 light bands from the axillar region to the leg base, juveniles and females with one or two V-shaped light bands on the occiput and nape (bands sometimes hardly visible in older specimens). Natural History Notes: No details, probably as for Hypsilurus macrolepis. SOLOMONS TREE DRAGON Hypsilurus macrolepis (Peters, 1872) Figs. 5, 6, 7 Distribution: Endemic to Solomon Islands: Makira, Ugi, Olu Malau and Santa Ana. Type locality: Solomon Islands. Description: Average SVL 120mm, tail about 450% (males) or 350% (females) of SVL. Head large and angular, slightly concave above. Canthus rostralis distinct. Body strongly compressed laterally. Adpressed hindlimb reaches the level of the snout. Head covered with small keeled scales decreasing in size posteriorly. A single row of enlarged scales runs from the mental, along the angle of the lower jaw to below the ear where they disperse into a cluster of enlarged, slightly keeled scales. Supralabials 10 to 12. Tympanum distinct, equal in size to the eye. Gular region with smooth scales anteriorly, becoming keeled or spinose posteriorly. A moderate gular fold is present. A well-developed (more so in males) nuchal crest is present, subcontinuous with the smaller dorsal crest. The dorsal crest continues onto the tail where it forms a slight caudal crest on the proximal third of the tail. The body scales are all strongly keeled, largest ventrally. Midbody scales in 75 to 110 rows. The caudal scales are larger than the body scales, all strongly keeled and slightly larger on the underside of the tail. No femoral or precloacal pores. Colour: Males: Dorsal ground colour grey-green to grey-brown. The posterior third of the body and proximal third of the tail are often a bright, iridescent mauve. A series of ill-defined light transverse bars is present on the posterior half of the body and continuing onto the tail for its full length. The limbs are the same ground colour as the body though they may be AGAMIDS, Family AGAMIDAE

25

mottled with lighter and darker spots. The throat is cream to light or dark grey or brick-red and in many individuals it is marked with yellow and orange and black spots. The ventral area is cream to yellowish without other markings. Females: Uniformly bright green above and yellow or yellow-green ventrally. The sides of the body may have a peppering of fine black spots. Juveniles (male and female) are bright green though the males are transversely banded with darker bars. Natural History Notes: This agamid is quite common throughout its range in the eastern Solomons. These lizards are almost completely arboreal though they will occasionally forage on the ground. They are very active diurnally and while they tend to favour the denser forest areas, they will also enter cleared areas at the edge of the forest as well as domestic gardens in villages and towns where there is ample vegetation. They move rapidly through the trees by a series of leaps from branch to branch. Like many arboreal animals they will move around on a tree so that the trunk is always between themselves and an observer. Females tend to be less active than males and seem to rely more on their cryptic colouration for protection. Food consists mainly of insects, particularly ants, spiders and very occasionally smaller lizards. Small amounts of flowers and fruits are also eaten. Unlike many agamids, H. macrolepis do not often bask in the sun for lengthy periods. They may be active in sunlit areas but they also spend long periods feeding and foraging in the shade. I have also seen them active during heavy rain. Like many agamids, male H. macrolepis tend to be territorial, actively defending their territories from intruding males. I have not observed courtship behaviour preliminary to mating in H. macrolepis. When mating occurs the female is seized and held in the jaws of the male and copulation takes place immediately. Two (very rarely 3) eggs are laid in rotting timber or in moist humus on the ground and the hatchling lizards emerge in about 6 to 8 weeks.

GECKOS, Family GEKKONIDAE A very large and diverse family of lizards which are distributed in tropical and temperate regions all around the world. Typically geckos are small to moderate sized lizards with pentadactyl limbs and with small granular scales covering their bodies. The head and body are usually flattened and the tongue is broad with a slight notch at its tip. Nearly all geckos have no

26

GECKOS, Family GEKKONIDAE

movable eyelids; the eye is covered with a modified transparent scale. The pupil is vertically elliptic in most species. An external ear opening is usually present and many species have femoral and precloacal pores. Their digits may be clawed or one or more digits may be clawless. Many species have modified subdigital lamellae which enable them to cling to vertical surfaces or to move upside down on the underside of branches and ceilings. Geckos are among the very few reptiles that have some ability to vocalize. The voluntary shedding of the tail (autotomy) is a common feature of most geckos. The regenerated tail usually differs from the original in scalation and colour and sometimes also in shape. Virtually all geckos are oviparous, tropical species typically laying 2 eggs at a time. Geckos may be either terrestrial or arboreal though almost all are nocturnal. They feed mostly on arthropods, though large geckos will eat hatchlings of their own and other species. The colours of most geckos change from day to night, generally the colouration during the day is markedly darker than at night when the lizard is active.

Subdigital lamellae (divided) Terminal lamella Claw Webbing

Digital expansion Underside of a gecko foot (Hemidactylus frenatus)

KEY TO THE GENERA OF SOLOMON ISLANDS GECKOS 1. All digits strongly clawed, digits not expanded distally: 2 – Four or 5 digits clawed, digits expanded distally: 3 GECKOS, Family GEKKONIDAE

27

2. A distinct lateral fold along the lower body between the fore- and hindlimbs: Cyrtodactylus – No lateral fold on the body: Nactus 3. Fifth digit moderately to well developed: 4 – Fifth digit rudimentary: Hemiphyllodactylus 4. Claw arising from within the extremity of the digital expansion: 5 – Claw arising from the extremity of the digital expansion, fifth digit clawless or with a minute claw: Lepidodactylus 5. Fifth digit clawless, body and tail slender, head large, unregenerated tail with a series of annulate scales: Gekko – Fifth digit with a minute claw, head and body moderate, tail depressed at its base and not noticeably slender: 6 6. Digits moderately dilated, fifth digit with a free terminal phalanx: Hemidactylus – Digits strongly dilated, fifth digit without a free terminal phalanx: Gehyra

Genus Cyrtodactylus Gray, 1827 Digits compressed, thin, strongly clawed. Claw between two scales. Enlarged postmental scales present. Femoral and precloacal present in males. KEY TO THE SPECIES OF SOLOMON ISLANDS Cyrtodactylus – Average SVL more than 100mm, underside of tail with transversely enlarged scales: louisiadensis – Average SVL less than 100mm, underside of tail without enlarged scales: biordinis

28

GECKOS, Family GEKKONIDAE

BANDED PALM GECKO Cyrtodactylus biordinis Brown and McCoy, 1980 Fig. 8 Distribution: Endemic to the Solomon Islands: Guadalcanal. Type locality: Mt Austen, Guadalcanal, Solomon Islands. Description: Average SVL 90mm, tail about 100% of SVL. Head large, slightly depressed, covered with small conical scales. Rostral with a slight median groove. Nostrils bordered by the rostral, first labial, a single large anterior circumnasal and 2 to 3 small posterior circumnasals which are often indistinguishable from the surrounding scales. Dorsally covered with very small conical scales with a series of enlarged conical granules forming more or less regular longitudinal rows. A distinct lateral fold is present. Tail slender, round in cross section, covered with subimbricate scales, no transversely enlarged scales present on its underside. Males with a double parallel row of femoral pores, usually separated from the single precloacal pore series, 60 or more pores in the femoral and precloacal series (not counting each pore in the doubled femoral series). Digits slender, compressed distally, strongly clawed. Colour: Dorsal ground colour light brown or grey-brown. A thick black band runs from the eye, along the neck in a dorsolateral position, usually joining the line on the opposite side of the body at a point slightly posterior to the forelimbs. A single large dark spot is present on the nape of the neck, 1 to 4 smaller spots are usually present on the occiput. 2 to 3 broad, unevenly margined dark bands are present dorsally, these bands edged posteriorly with white. The tail is banded black and white for the proximal third of its length and black for the distal portion except for the tip which is grey during the day and white at night when the gecko is active. Ventrally grey to grey-white. Natural History Notes: This fairly common arboreal gecko shelters by day in tree hollows in forested areas. At night it moves about on the trunks and branches of the smaller trees and the vines in the forest understorey, foraging for its insect food. It does not appear to inhabit or move about nocturnally on the larger bare-trunked forest trees to any extent. On such trees it tends to be replaced by Cyrtodactylus louisiadensis, a related species with which it is sympatric. When moving, C. biordinis often holds its tail arched above its back. Its 2 eggs are laid in the detritus in tree hollows and possibly also in ground debris at the bases of trees. GECKOS, Family GEKKONIDAE

29

RING-TAILED GECKO Cyrtodactylus louisiadensis (De Vis, 1892) Figs. 9, 10, 11 Fred Kraus (pers. comm.) believes the Solomons populations of this gecko are not referrable to true C. louisiadensis. Distribution: New Guinea and adjacent archipelagos, Cape York in Australia, Solomon Islands. In the Solomons: Bougainville, Shortland, New Georgia, Isabel, Guadalcanal, Malaita. Type locality: Tagula Island (Louisiade Archipelago), Papua New Guinea. Description: Average SVL 130mm, tail about 120% of SVL. Head large, wedge-shaped, slightly depressed and covered with very small conical scales with larger granules present on the supraocular and occipital regions. Rostral quadrangular, a little wider than deep with a short median groove posteriorly. Mental as wide as the rostral, followed by a pair of smaller postmentals. Nostril bordered by the rostral, first labial, a large anterior circumnasal and 3 to 4 small posterior circumnasals which are often indistinguishable from the surrounding scales. Dorsally covered with very small conical granules with a regular series of enlarged conical scales. Ventrally with smooth imbricate scales. A strong lateral fold is present. Tail round in cross section, slightly flattened at its base with transversely enlarged scales on its underside. Males with 40 to 70 femoral and precloacal pores. All digits slender, slightly compressed distally and strongly clawed. Colour: Dorsally the body is light yellowish brown to medium brown with several (usually 4) broad darker crossbands, each band with narrow lighter borders. A similar V-shaped darker band runs from eye to eye across the occiput. The tail is banded with dark brown and white. At night when the gecko is active, the distal third to half of the tail is white. Ventrally grey to yellowish white. Natural History Notes: This large gecko is arboreal and prefers to inhabit the larger forest trees where it shelters in hollows or under loose bark during the day; the hollows in the strangler fig trees Ficus spp. are a favourite habitat of C. louisiadensis. At night it may very occasionally be seen on tree trunks a few metres above the ground or on the buttress roots of large rainforest trees, usually in a head-down position where it is presumably scanning the ground for arthropod prey. It is an active predator on insects, spiders and smaller geckos. This gecko is rarely seen,

30

GECKOS, Family GEKKONIDAE

though this may be a reflection of the fact that it tends to forage nocturnally in the crowns and upper branches of the larger forest trees and infrequently moves low on the trunk.

Genus Gehyra Gray, 1834 Digits strongly dilated with undivided or medially divided subdigital lamellae. All digits clawed (the fifth digit with a minute claw). The claw (not the minute fifth claw) arises from within the extremity of the digital expansion. Inner digits without free terminal phalanges. Femoral and precloacal pores present in males. KEY TO THE SPECIES OF SOLOMON ISLANDS Gehyra – Subdigital lamellae undivided: oceanica – Subdigital lamellae mostly divided: mutilata STUMP-TOED GECKO Gehyra mutilata (Wiegmann, 1835) Fig. 12 Distribution: Madagascar, Sri Lanka, southeast Asia, Japan, Philippines, Cocos (Keeling) Islands, Christmas Island, New Guinea, most of the islands of tropical Oceania, Mexico. In the Solomons it has been recorded from Bougainville, Guadalcanal (only recorded from houses at Marau at the eastern end of the island), Makira (only recorded from houses in the provincial capital, Kira Kira), Santa Ana (only recorded from village houses) and Graciosa Bay on Santa Cruz Island. Given such disparate and localised populations, it would have to be assumed that the distribution of this species in the Solomons is a result of human agency. Type locality: Manila, Philippines. Description: Average SVL 55mm, tail about 100% of SVL. Head moderate, slightly depressed, covered with small conical granules. Body with a slight lateral fold. Rostral quadrangular, wider than deep. Mental pentagonal, followed by a pair of much larger postmentals and several pairs of enlarged chinshields. Nostril bordered by the rostral, first labial, a single large anterior circumnasal and usually two small posterior circumnasals. Dorsally the body is covered with smooth conical scales which increase in size laterally. Tail quite depressed and fairly wide at its GECKOS, Family GEKKONIDAE

31

base though often a little constricted at its point of attachment to the body, covered with slightly spinose scales above, giving the tail a noticeably serrate edge. Underside of the tail with large transverse scales. Ventrally with smooth imbricate scales. Digits webbed at their bases and strongly dilated with 8 to 10 lamellae under the median toes, the terminal lamellae are undivided, most of the rest are divided. Males with 32 to 39 femoral and precloacal pores. Colour: Head and dorsum grey to light brown with a peppering of lighter and darker spots and blotches. At night when the gecko is active, the colour is a uniform pinkish-white. Ventrally light yellow or cream. Natural History Notes: An arboreal gecko often occurring as a human commensal. In the Marau area at the eastern end of Guadalcanal, I have seen G. mutilata at night on interior and exterior walls of houses together with the common house gecko Hemidactylus frenatus, and the occasional oceanic gecko Gehyra oceanica, and mourning gecko Lepidodactylus lugubris. Earlier literature does not list this species as occurring in the Solomons; it is probably a recent introduction from elsewhere in the Pacific, having been brought in by shipping as eggs or mature lizards. The specific name of mutilata refers to this gecko’s tendency to shed large portions of skin if it is grasped even lightly. This is a means of avoiding predation analogous to the tail shedding of many skinks and geckos, and pieces of skin lost are soon regrown by the gecko. OCEANIC GECKO Gehyra oceanica (Lesson, 1830) Fig. 13 Distribution: Indo-Australian archipelago, New Guinea, most of the islands of tropical Oceania. In the Solomons: Bougainville, Shortland, Mono, Choiseul, Rob Roy, New Georgia, Tetepare, Vangunu, Isabel, Guadalcanal, Tulagi, Ngela, Malaita, Ontong Java, Makira, Ugi, Olu Malau, Bellona, Santa Cruz, Taumako (Duff Islands), Reef Islands, Utupua. It is likely that this wide-ranging species occurs on most islands of the Solomons. Type locality: Tahiti and Bora Bora. Description: Average SVL 90mm, tail about 100% of SVL. Head moderate, slightly depressed. Body stout with an indistinct lateral fold. Hindlimbs with a distinct fold along their posterior edges. Nostril bordered

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GECKOS, Family GEKKONIDAE

by the rostral, first labial, a single anterior circumnasal and 3 to 4 smaller posterior circumnasals. Triangular mental followed by a row of smaller postmentals. Dorsally covered with small smooth conical scales. Ventrally with smooth imbricate scales. The tail is covered with subimbricate scales, largest on its underside. All digits clawed, the fifth digit with a minute claw. Claws (except the fifth claw) arise from within the extremity of the digital expansion. Fifth digit without a free terminal phalanx. Digits all strongly dilated with an undivided series of 11 to 18 lamellae under the median toes. Males with 26 to 42 femoral and precloacal pores. Colour: Ground colour light to darker brown with irregular lighter and darker flecking over the dorsum, limbs and tail. The colour fades at night to a pale creamy white with few distinct markings. Ventrally cream to yellow. The underside of the tail is often bright yellow or orange. Natural History Notes: A common arboreal species, sometimes found in human habitations, though it is not generally a human commensal. In forest areas it is often found on the larger trees, resting in an exposed position on the trunk during the day, relying on its cryptic colouration for protection. It is also a common inhabitant of the crowns of coconut and sago palms. It feeds on insects and spiders and occasionally preys on smaller geckos. It also licks nectar from flowering plants and will feed on very ripe fruit - such as mango, banana and pawpaw - on an opportune basis, particularly where the fruit has already been partially eaten by birds or flying foxes. It lays 2 eggs under loose bark, in tree hollows or occasionally in the ceilings of houses. Communal egg-laying is not uncommon in this species, where up to 20 or more eggs may be laid at the same site. At Boromole on Ngela Island I found a communal egg site with more than a dozen eggs under rotting sago palm leaves on the ground. Like Gehyra mutilata, G. oceanica will shed pieces of its skin if it is grasped.

Genus Gekko Laurenti, 1768 Digits strongly dilated with a series of undivided subdigital lamellae. All digits except the fifth, clawed. Claws arise from within the extremity of the digital expansion. Femoral and precloacal pores present in males.

GECKOS, Family GEKKONIDAE

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SAGO GECKO Gekko vittatus Houttyn, 1782 Figs. 14, 15 Distribution: India, Indonesia, Palau (Caroline Islands), New Guinea and adjacent island archipelagos, Solomon Islands. In the Solomons: Bougainville, Mono, Vella lavella, New Georgia, Gizo, Vangunu, Guadalcanal, Ngela, Makira, Olu Malau, Santa Ana, Santa Catalina, Rennell, Bellona, Santa Cruz, Utupua. Type locality: India. Description: Average SVL 140mm, tail about 120% SVL. Head very large, distinctly depressed. Body elongate with a lateral fold. Limbs well developed, slender. Tail round in cross section, prehensile, slender, with a series of annulate scales. Rostral quadrangular, wider than deep. Nostril bordered by the rostral, first labial, a single large anterior circumnasal and 2 to 3 smaller posterior circumnasals. Triangular mental followed by numerous slightly enlarged scales. Head and dorsum covered with small, smooth conical scales. A regular series of enlarged conical tubercles is present on the occipital region of the head, dorsum and limbs. Ventrally with smooth imbricate scales. Digits strongly dilated, fifth digit clawless, claws arising from within the extremity of the digital expansion. 18 to 20 lamellae in an undivided series under the median toes. Males with 14 to 58 femoral and precloacal pores. Colour: Ground colour variable, usually light or dark brown or grey. A narrow white vertebral stripe is usually present in most populations, running from the base of the tail and onto the neck where it forks, each branch fading out on the occipital region. The tail is usually marked with broad whitish bands. Ventrally white to greyish yellow. Populations in the eastern Solomons lack the vertebral stripe (though it is usually present in juveniles) and are marked with irregular dark brown and white blotches on the head, body and tail. Natural History Notes: A common arboreal species. During the day this gecko often remains motionless on exposed tree trunks or along thin vines and branches, relying on its cryptic colouration for protection as does Gehyra oceanica, a species with which it is often found. Another favoured diurnal retreat is among the dead, trailing leaves of banana and pandanus trees. At night it is an active voracious predator on smaller geckos, insects and spiders. It moves swiftly through the trees with the aid of its prehensile

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GECKOS, Family GEKKONIDAE

tail and also by leaping from branch to branch over distances of up to 1m. It is a common house gecko in parts of the eastern Solomons. It lays its 2 eggs, which adhere to each other and to the substrate on which they are laid, under loose bark or on the underside of leaves. I have observed community egg-laying in this species in pandanus trees where up to 100 eggs and remains of eggs representing several generations may be present among the dead leaves of a single tree.

Genus Hemidactylus Gray, 1825 Digits free or webbed, moderately dilated with a series of medially divided subdigital lamellae. All digits clawed, fifth digit with a very small claw. The digits arise from within the extremity of the digital expansion. Fifth digit with a free terminal phalanx. Femoral and precloacal pores present in males. COMMON HOUSE GECKO Hemidactylus frenatus Duméril and Bibron, 1836 Fig. 16 Distribution: Southern and eastern Africa, Madagascar, Mauritius, Sri Lanka, India, southern China, Japan, Philippines, southeast Asia, Cocos (Keeling) Islands, Christmas Island, New Guinea, northern Australia, Oceania, Mexico, parts of the southern United States, St Helena. In the Solomons: Bougainville, Shortland, Taro Island (in Choiseul Bay), Gizo, New Georgia (Munda), Isabel (Buala), Guadalcanal (Honiara, Marau), Tulagi, Malaita (Auki), Makira (Kira Kira), Santa Cruz (Lata). Type locality: Java. Description: Average SVL 55mm, tail about 120% of SVL. Head moderate. Rostral quadrangular with a distinct median groove. Nostril bordered by the rostral, first labial, a single large anterior circumnasal and 2 small posterior circumnasals. Mental triangular, as wide as deep, followed by a pair of enlarged postmentals. Dorsally covered with small, smooth conical scales. There are enlarged conical granules at regular intervals along the body dorsally and laterally. Ventrally with smooth imbricate scales. Tail prehensile, flattened at its base, covered with smooth imbricate scales and a regular series of enlarged spinose scales forming annuli. The underside of the tail has enlarged transverse scales. All digits clawed, the fifth digit with a very small claw. The claws arise from within GECKOS, Family GEKKONIDAE

35

the extremity of the digital expansion. Digits moderately dilated with a series of 7 to 10 divided lamellae under the median toes. Males with 20 to 40 femoral and precloacal pores. Colour: Grey to whitish above with a peppering of darker flecks that may form regular patterning in some specimens. These markings are only evident during the day when the gecko is at rest, at night the colour is a uniform pinkish white though the tail is often a light orange-pink. Ventrally whitish. Natural History Notes: This species is only found as a commensal of man wherever it occurs in the Solomons, its presence indicated by its distinctive chirping call. Regular electric lighting is a factor in this gecko’s establishing breeding populations in buildings. In the evenings it is often abundant around interior and exterior house lights, feeding on the insects that are attracted to these lights. In this situation it is not unusual to see up to a dozen or more geckos gathered around a light. Bustard (1970), studying this species in New Guinea, found that the greatest activity is reached in the hour following sunset and then sharply decreases until a minor increase again at dawn. Earlier workers invariably did not include this species in the Solomons reptile fauna, most probably indicating that it is a relatively recent introduction from elsewhere. This likelihood becomes more probable when one considers the affinity of the gecko for dwelling in houses and laying its 2 eggs not only in the ceilings, but also in building materials for example that may be shipped from island to island. It is possible that H. frenatus was introduced during World War II when large amounts of military equipment were brought into the Solomons from other Pacific countries. At the present time, H. frenatus appears to be restricted to the larger towns in the Solomons. No doubt with increasing movement of people and material between the islands of the group, and the extension of electrical lighting into more rural communities, H. frenatus will eventually become more widespread.

Genus Hemiphyllodactylus Bleeker, 1860 Digits dilated, without webbing. Fifth digit rudimentary, clawless, other digits clawed with free terminal phalanges, the claws arising from within the extremity of the digital expansion. Femoral and precloacal pores are present.

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GECKOS, Family GEKKONIDAE

Hemiphyllodactylus typus Bleeker, 1860 Fig. 17 Distribution: Sri Lanka, southern India, Philippines, Taiwan, southeast Asia, Indonesia, New Guinea, Solomon Islands, much of Oceania. Apparently introduced into Hawai’i by early Polynesian voyagers (Kluge, 1966). In the Solomons it has been recorded from Bougainville and Guadalcanal. Type locality: Java. Description: Average SVL 40mm, tail about 100% of SVL. Head moderate, longer than wide. Body elongated, slender. Limbs rather short. Rostral quadrangular. Nostril bordered by the rostral, first labial, a single large anterior circumnasal and 2 small posterior circumnasals. Mental triangular, followed by 1 to 3 rows of enlarged scales. 8 to 12 supralabials. Body covered with small granular scales without larger tubercles. Tail round in cross section, slender. Fifth digit rudimentary, clawless and with 4 to 5 undivided lamellae. Other digits clawed, the claws arising from within the extremity of the digital expansion. Fourth digit with 4 to 5 yshaped lamellae, usually medially divided. 6 to 11 precloacal pores and 8 to 13 femoral pores. Colour: Dorsally light brown to reddish brown with scattered darker flecks. A dark stripe runs from the snout, through the eye and onto the neck. A series of round lighter spots is often present dorsolaterally. Ventrally whitish speckled with brown. At night when the gecko is active, the colour is a pale pinkish white. Natural History Notes: This is an all-female species that reproduces by parthenogenesis. It is essentially an arboreal gecko, sheltering by day in the leaf axils of ferns and palms and under the bark of trees. In its range outside the Solomons it has been reported as occasionally living in human habitations. The lizard illustrated (fig. 17) I found at night on the exterior wall of my house in Honiara on Guadalcanal. A pair of gecko eggs I found adhering to the underside of a Heliconia leaf in a forested area near the coast on Guadalcanal proved to be this species on hatching. This gecko is uncommon throughout most of its range and apparently very rare in the Solomons.

GECKOS, Family GEKKONIDAE

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Genus Lepidodactylus Fitzinger, 1843 Digits moderately dilated with a partially divided or an undivided series of subdigital lamellae. The claw arises from the extremity of the digital expansion. Fifth digit clawless or with a very small claw. Males with femoral and precloacal pores. KEY TO THE SPECIES OF SOLOMON ISLANDS Lepidodactylus 1. Terminal lamellae divided: 2 – Terminal lamellae undivided: 3 2. Tail more or less distinctly flattened with spinose scales along its edges forming a lateral fringe, digits with at least some degree of webbing, all known Solomons populations are parthenogenetic females: lugubris – Tail only slightly flattened, spinose scales along its edges poorly developed and not forming a distinct fringe, digits without webbing, Solomons populations are bisexual: woodfordi 3. All lamellae undivided: 4 – Several distal lamellae divided: 5 4. 9 to 11 lamellae under the fourth toe of the hind limb, 27 to 34 femoral and precloacal pores: mutahi – 18 lamellae under the fourth toe of the hind limb, 38 femoral and precloacal pores: flaviocularis 5. 11 to 15 lamellae under the fourth toe of the hind limb, 32 to 38 interorbital scales, 11 to 13 supralabials, males with 35 to 42 femoral and precloacal pores: guppyi – 11 lamellae under the fourth toe of the hind limb, 23 interorbital scales, 10 supralabials, 32 femoral and precloacal pores: shebae YELLOW-EYED GECKO Lepidodactylus flaviocularis Brown, McCoy and Rodda, 1992 Fig. 18 Distribution: Endemic to Solomon Islands: Guadalcanal. Type Locality: Mount Austen, Guadalcanal, Solomon Islands.

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GECKOS, Family GEKKONIDAE

Description: (of the holotype, an adult male) SVL 49mm, tail abbreviated. Digits moderately dilated with moderate webbing. 18 undivided lamellae under the fourth toe of the hind limb. Supralabials 12, the tenth below the centre of the eye. 38 precloacal and femoral pores. Nostril bordered by the rostral, first labial, a supranasal and 2 enlarged scales. Supranasals separated by 3 scales. Mental followed by several rows of only slightly enlarged scales. Hindlimbs with a strong fold on their posterior edge. The body and head are covered with very small, smooth conical scales with no enlarged tubercles. Colour: Light yellowish-tan with irregular darker wavy lines. The scales around the margin of the eye are bright yellow forming a distinctive orbital ring. Ventrally yellowish-cream, becoming bright yellow in the region of the vent. Natural History Notes: Known only from the male holotype collected in 1990 and a female specimen (fig. 18) I collected in 1978 (this specimen was forwarded to Walter Brown, but is apparently now lost). Both animals were found at night near the summit of Mount Austen (about 320m elevation) in secondary rainforest and both were found about 3m above the ground, the holotype on a thin vine and the female on the leaf axil of a small palm. The type locality on Mount Austen is now severely degraded by logging and subsistence gardening. GUPPY’S GECKO Lepidodactylus guppyi (Boulenger, 1884) Figs. 19, 20 Probably a composite species. Distribution: Bismarck Archipelago (Papua New Guinea), Solomon Islands. In the Solomons: Bougainville, Fauro, Kolombangara, New Georgia, Tetepare, Isabel, Guadalcanal, Ngela, Savo, Malaita, Makira, Olu Malau, Santa Cruz, Taumako (Duff Islands), Vanikoro. Type locality: Fauro Island, Solomon Islands. Description: Average SVL 45mm, tail about 120% of SVL. Head small. Body slender and elongate. Tail slender, round in cross section. Rostral quadrangular, wider than deep. Nostril bordered by the rostral, first labial and 3 to 4 small circumnasals. Supralabials 11 to 13. Mental followed by several rows of slightly enlarged chinshields. Head and body covered with small, smooth conical granules. Ventrally with smooth subimbricate scales. GECKOS, Family GEKKONIDAE

39

Digits moderately dilated, fifth digit clawless or with a minute claw, other digits clawed, the claw arising from the extremity of the digital expansion. Digits with a slight web at their bases. 11 to 15 lamellae under the fourth toe, terminal lamellae undivided, 2 to 4 subterminal lamellae divided. Males with 35 to 42 femoral and precloacal pores. Colour: Dorsally light to dark brown with a more or less regular series of lighter broad, wavy transverse bands edged anteriorly with black, these bands continuing onto the tail. An alternating series of larger black and whitish spots usually form a line in a ventrolateral position between the fore and hindlimbs. The head is marked irregularly with lighter and darker blotches. There is often a dark streak on the side of the head, passing from the snout, through the eye and onto the neck. Ventrally uniformly yellowish white, or with darker speckling. The colour and pattern is somewhat variable in this species. Natural History Notes: An uncommon arboreal species and little is known of its ecology. Its habits are probably similar to Lepidodactylus lugubris excepting that it does not enter human habitations to any extent, preferring to shelter diurnally under loose bark and very occasionally under ground debris. Like L. lugubris it also commonly shelters, and lays its 2 eggs, in the leaf axils and dead trailing leaves of pandanus trees. MOURNING GECKO Lepidodactylus lugubris (Duméril and Bibron, 1836) Fig. 21 Distribution: Sri Lanka, India, islands of the Indian Ocean, Philippines, Japan, Burma, Malaysia, Indonesia, Cocos (Keeling) Islands, Christmas Island, New Guinea and adjacent island archipelagos, northern Australia, most of tropical Oceania, Mexico, central America. In the Solomons: Bougainville, Shortland Islands, Choiseul, Vella lavella, Ranongga, New Georgia, Guadalcanal, Ngela, Malaita, Ontong Java, Rennell, Makira, Olu Malau, Santa Ana, Santa Catalina, Santa Cruz, Taumako (Duff Islands), Reef Islands, Vanikoro, Utupua, Tikopia. Type locality: Tahiti. Description: Average SVL 40mm, tail about 110% of SVL. Head small, body moderate. Tail stout, depressed, particularly at its base. The scales on the edge of the tail are enlarged and spinose and form a distinct lateral fringe. Nostril bordered by the rostral, first labial, a single large anterior circumnasal and 2 smaller posterior circumnasals. Mental followed by

40

GECKOS, Family GEKKONIDAE

several rows of enlarged chinshields. Supralabials 10 to 12. The digits are moderately dilated, the fifth digit with a minute claw. The claws arise from the extremity of the digital expansion. 11 to 15 lamellae under the fourth toe, the terminal and 3 or 4 distal lamellae are divided. Colour: Variable. The commonest colour form is pale fawn to grey above with a series of indistinct, often obscure, darker W-shaped markings along the centre of the back and onto the tail. There is usually a dark streak running from the snout, through the eye and onto the neck. Ventrally yellowish to white. Natural History Notes: A common small arboreal gecko, often occurring as a human commensal. Its normal diurnal retreat in forest areas is under loose bark or amongst dead trailing foliage. It is also a common inhabitant of the leaf axils of pandanus and coconut palms. It lays 2 eggs, which adhere to each other and to the substrate on which they are laid, under bark and leaves and also in the ceilings of houses. Community egg laying occurs in this species; many individuals of L. lugubris may share a single pandanus tree for oviposition. All known Solomons populations of L. lugubris consist entirely of females and reproduction is by parthenogenesis. Pairs of these geckos often indulge in pseudo-copulation with one of the pair assuming the role of the male lizard, biting and holding the other by the neck and wrapping its tail around to bring their cloacas into contact. It has been theorised that this pseudo-copulation may stimulate the unfertilised ovum to begin cell division. Lepidodactylus lugubris is not a particularly active species when compared with many other geckos, when feeding it tends to wait for its insect prey to come within reach rather than actively making pursuit for any distance. Given the opportunity L. lugubris feeds on food left unattended in houses, particularly sweet foods. I have seen eight geckos gathered around a open tin of jam, eagerly licking the contents. I have also often seen them drinking the dregs in cups of sweetened tea and coffee. MUTAHI SCALY-TOED GECKO Lepidodactylus mutahi Brown and Parker, 1977 Distribution: Endemic to the Solomon Islands: Bougainville. Type locality: Mutahi, Bougainville Island. Description: Average SVL 45mm. Body moderately depressed, slightly elongate. Tail subcylindrical. Nostril bordered by the rostral, first labial, a GECKOS, Family GEKKONIDAE

41

single anterior circumnasal and 2 smaller posterior circumnasals. Supralabials 10 to 12, infralabials 9 to 11. Mental triangular, followed by several rows of enlarged chinshields. Body covered with small granular scales without larger tubercles. Digits moderately dilated, slight webbing between the toes. All digits except the fifth finger and toe, clawed, the claws arising from the extremity of the digital expansion. 9 to 12 undivided lamellae beneath the fourth toe. Males with 27 to 34 femoral and precloacal pores. Colour: Dorsally pale yellow-brown to dark red-brown, often with a light stripe running from the eye to the ear. The tail has orange patches and the whole body is often marked with darker blotches or bars. Natural History Notes: Brown and Parker (1977) report this species as being most abundant in pandanus in permanent coastal swamps, but that it also occurs in banana trees and newly cleared garden areas. GUADALCANAL SCALY-TOED GECKO Lepidodactylus shebae (Brown and Tanner, 1949) Distribution: Known only from the Solomon Islands: Guadalcanal. Type locality: lower Lungga River area, Guadalcanal Island, Solomon Islands. Description: (Adapted from Brown and Tanner 1949) SVL 36mm, tail 29mm. Head about 1.5x as long as broad. Body slender. Tail round in cross section, slender. Limbs well developed. Rostral more than 2x as wide as deep, with a median groove dorsally. Nostril bordered by the rostral, first labial and 3 circumnasals. Supralabials 10, infralabials 9. Mental pentagonal, followed by 2 rows of chinshields and 2 rows of slightly enlarged scales. 23 interorbital scales. The head and body are covered with small granular scales. Ventrally with larger imbricate scales. Digits long, evenly dilated with partial webbing. Fifth toe clawless. The claws on the other digits are contained in a sheath. 11 lamellae under the fourth toe, terminal lamellae undivided, 2 or 3 distal lamellae divided. 32 femoral and precloacal pores. Colour: Reddish brown dorsally with lighter mottling laterally. A narrow broken band runs from beneath the eye and onto the neck. A series of 3 small brown blotches is present in a dorsolateral position on each side of the body from the neck to a little posterior to the forelimb. The tail has distinct transverse bands dorsally. Natural History Notes: This species is known only from the holotype collected in the “...lower Lungga River area” on Guadalcanal in 1944

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GECKOS, Family GEKKONIDAE

(Brown and Tanner, 1949). No known habitat or other ecological data exist for the unique specimen. WOODFORD’S SCALY-TOED GECKO Lepidodactylus woodfordi Boulenger, 1887 Fig. 22 Boulenger described this species from a single specimen which he erroneously allied to L. guppyi whereas it is very closely related to L. lugubris. The validity of L. woodfordi has been questioned by Brown and Parker (1977). However, L. woodfordi is bisexual and this would distinguish it from the widespread parthenogenetic forms of L. lugubris. The specimen illustrated (fig. 22) is a male. Distribution: Papua New Guinea, Solomon Islands. Type locality: Fauro Island, Solomon Islands Description: The following is Boulenger’s original description: “Closely allied to Lepidodactylus guppyi. Digits without distinct web. Tail a little depressed, rounded. Femoral and precloacal pores 25 altogether. Grey above with zigzag black cross bands, six between the nape and base of the tail; a black streak from the nostril to the neck passing through the eye and above the ear; lower surfaces whitish.” Boulenger further gives the SVL as 40 mm and the tail as 38 mm. Natural History Notes: Geckos that I believe to be referrable to L. woodfordi in the sense of Boulenger (1887b) I have seen on the ceilings and walls of houses at Korovou on Shortland Island (adjacent to Fauro Island), feeding on small insects attracted to the house lights.

Genus Nactus Kluge, 1983 Digits long and slender and strongly clawed. Males with precloacal pores. SOLOMONS SLENDER-TOED GECKO Nactus multicarinatus (Günther, 1872) Fig. 23 This species is a bisexual relative of the wider ranging Nactus pelagicus, which consists of unisexual populations throughout much of Oceania. GECKOS, Family GEKKONIDAE

43

Distribution: Solomon Islands, Vanuatu. In the Solomons: virtually every island of the group from Bougainville to the Eastern Outer Islands. Type locality: Vanuatu. Description: Average SVL 60mm, tail about 100% of SVL. Head large, slightly depressed, covered with small conical scales. Rostral with a median groove. Nostril bordered by rostral, first labial, a single large anterior circumnasal and 3 to 4 small posterior circumnasals that are often indistinguishable from surrounding scales. Dorsally covered with small multi-keeled conical scales with a series of enlarged, multi-keeled conical tubercles forming regular longitudinal rows, there are usually 16 to 18 of these longitudinal rows at midbody. Tail slender, evenly tapered, round in cross section, covered with small, slightly spinose scales. Males with 6 to 10 precloacal pores. Digits long, slender, strongly clawed. Colour: Dorsally grey-brown with irregular darker wavy transverse bands edged posteriorly with white, this banding continued onto the tail. Ventrally grey to almost white. The lips are usually barred with white. At night when the gecko is active, the dorsal pattern is often obscure and the whole body is light grey in colour. Natural History Notes: A very common terrestrial gecko. It occurs in both forested areas and also in plantations and domestic gardens in towns around the Solomons. It shelters diurnally under any suitable ground debris such as rotting fallen timber and amongst rocks and in crevices in the root systems of large trees. It is also a very common inhabitant of piles of coconut husks in plantations. Active and agile at night, when pursuing prey or escaping from a potential predator it will often move in a series of rapid short leaps, running for several steps between jumps. Although it forages mainly on the ground, it also climbs a little distance up the trunks of trees at night in search of its insect prey. It lays its two eggs in and under ground debris or in hollows around the roots of trees.

SKINKS, Family SCINCIDAE This family contains a large assemblage of lizards differing greatly in size, body form and ecology, and in the Solomons they represent the majority of reptile species. Skinks occur virtually all over the world, from the large continental land masses to the far-flung islands of the Pacific, reaching

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SKINKS, Family SCINCIDAE

their greatest diversity and abundance in the tropics. Although most species are terrestrial, many are largely arboreal and others are fossorial in their habits. Although they are mostly diurnal lizards, a number are crepuscular or nocturnal, particularly the burrowing species. The lower eyelid of skinks may be scaly or a transparent palpebral disc may be present. Other forms have an immovable transparent spectacle covering the eye similar to the geckos. Also like the geckos, many skinks have the ability to autotomise their tails, subsequently regenerating them. The head shields of skinks are more or less symmetrical and the tongue is short and broad. The smaller species feed on arthropods and some of the larger forms are partly or completely herbivorous. Reproduction in skinks is by egglaying or by live birth.

Parietal Interparietal Supraoculars

Temporals Supralabials

Rostral Tympanum

Nuchals

Loreal

Frontonasal Prefrontal Frontal Frontoparietal

Nasal Infralabials

Head scalation (dorsal and lateral) of a skink (Lamprolepis smaragdina)

Methods of counting reptile midbody scale rows. The diagram above represents the scalation of a snake, the same method is used to count lizard midbody scales SKINKS, Family SCINCIDAE

45

KEY TO THE GENERA OF SOLOMON ISLANDS SKINKS 1. Forelimbs with five digits: 2 – Forelimbs with four digits: Carlia 2. Median internuchal scale present, prefrontals absent, body scales smooth: Geomyersia – Median internuchal scale absent, prefrontals present or if absent then body scales carinate or spinose: 3 3. Lower eyelid movable, scaly or with a palpebral disc: 4 – Eye covered with an immovable transparent spectacle: Cryptoblepharus 4. Lower eyelid scaly: 7 – Lower eyelid with a palpebral disc: 5 5. Supranasals present, 30 or more lamellae under the fourth toe of the hindlimb: Emoia – Supranasals absent, fewer than 30 lamellae under the fourth toe of the hindlimb: 6 6. Proximal lamellae expanded, uniformly pale lime green or olive green dorsally: Prasinohaema – Lamellae not expanded, dorsally brownish with a light vertebral stripe: Lipinia 7. Body scales smooth: 8 – Body scales carinate or spinose: Tribolonotus 8. Body elongate, limbs poorly to moderately well developed, dorsally brown with lighter and/or darker markings: 10 – Body not noticeably elongate, limbs well developed, dorsal ground colour pale green to olive green, bronze green or bright green: 9 9. Very large size (average SVL 350 mm), tail slender, prehensile, parietals widely separated: Corucia

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SKINKS, Family SCINCIDAE

– Moderate size (average SVL 90 mm), tail evenly tapered, not prehensile, parietals in contact: Lamprolepis 10.Auricular lobules present, prefrontals in contact or separated, supranasals present: Eugongylus – Auricular lobules absent, prefrontals in contact or separated, supranasals absent or if present then head shields partially fragmented: Sphenomorphus

Genus Carlia Gray, 1845 Small terrestrial, diurnal lizards. Forelimbs with 4 digits, hindlimbs with 5 digits. Frontoparietals fused into a single shield. Lower eyelid with a palpebral disc. Parietals in contact. Oviparous. Because of the iridescent sheen on the scales of these lizards, they are commonly called rainbow skinks. MYS’ RAINBOW SKINK Carlia mysi Zug, 2004 Fig. 24 Distribution: North coast of New Guinea from Manam Island eastward to Cape Vogel, Bismarck Archipelago, Solomon Islands. In the Solomons: Bougainville, Shortland, Faisi Island (Shortland group). Type locality: Gusika, Morobe Province, Papua New Guinea. Description: Average SVL 55mm, tail about 150% of SVL. Head moderate. Body a little stout. Limbs well developed, overlapping when adpressed. Forelimbs with 4 digits, hindlimbs with 5 digits. Supranasals absent. Prefrontals usually separated. Frontal a little longer than wide, longer than its distance from the snout, in contact with the first and second of 4 (rarely 5) supraoculars. Lower eyelid with a palpebral disc. Frontoparietals fused into a single shield. A small interparietal is present. Supralabials 7 (rarely 6 or 8). Enlarged nuchal scales present. The ear opening has several small overlapping lobules at its anterior edge. The dorsal and lateral body scales are weakly carinate. Midbody scales in 28 to 36 rows, 23 to 34 lamellae under the fourth toe. Colour: Dorsal ground colour a uniform light to dark brown. Juveniles sometimes with white dorsolateral and midlateral stripes bordering a SKINKS, Family SCINCIDAE

47

darker lateral stripe. This pattern is occasionally found in adult females and less frequently in adult males. Ventrally yellowish white. Body with an iridescent sheen. Natural History Notes: The occurrence of this species on Bougainville is apparently limited to the major towns and their immediate vicinities. It is possible that it has been introduced into Bougainville (and from there to the Shortlands) from mainland New Guinea or New Britain where it is also common in and around towns. In 1983, on Faisi Island in the Shortland group, I found Carlia mysi occurring very commonly in, and apparently strictly limited to, the overgrown grassy areas adjacent to the site of a preWorld War II trade store, long demolished. Where the grassy tracts at this site merged into open scrub, C. mysi was immediately replaced by Emoia cyanura. As I have not returned to Faisi since that time I do not know the current status of that isolated population of Carlia mysi. It is an active terrestrial skink and its preferred habitat, at least on Bougainville, is amongst grass tussocks in open and semi-cleared areas.

Genus Corucia Gray, 1855 A monotypic genus. Head large, limbs well developed and strongly clawed, tail prehensile. Lower eyelid scaly. No supranasals. Prefrontals in contact or narrowly separated. Parietals widely separated. PREHENSILE-TAILED SKINK Corucia zebrata Gray, 1855 Figs. 25, 26, 27 Köhler (1997) has referred the Bougainville and Buka populations to the subspecies Corucia zebrata alfredschmidti based to a large extent on the colour and pattern of live animals. On the basis of his description, this form also occurs on Shortland Island and is illustrated in fig. 25. It is worth noting that taxonomic assessments of Corucia populations based on captive specimens outside the Solomons should be viewed with some reservations. The provenance of such specimens (if obtained through the pet trade) cannot be regarded with any certainty, and captive bred populations of Corucia may well be the descendants of original stock from widely separated islands.

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SKINKS, Family SCINCIDAE

Distribution: Endemic to the Solomon Islands archipelago: Bougainville, Shortland Islands, Choiseul, Vella lavella, New Georgia, Tetepare, Vangunu, Isabel, Guadalcanal, Ngela, Malaita, Makira, Ugi, Santa Ana. Type locality: Makira Island (San Cristobal), Solomon Islands. Description: Average SVL 350mm, tail about 120% of SVL. Head large, wedge-shaped, snout short. Body quite robust. Limbs well developed, digits strongly clawed. Tail slender, prehensile. The frontonasal is the largest head shield, prefrontals in contact or separated, sometimes by a small median scale, frontal about as wide as long, frontoparietals distinct, lower eyelid scaly, parietals widely separated, bordered by one or more pairs of enlarged temporals, nuchals usually enlarged. The head shields are often irregular and subject to variation. Midbody scales in 35 to 40 rows, 19 to 22 lamellae under the fourth toe. Colour: Dorsal ground colour variable, ranging from pale olive green to grey-green to khaki with lighter and darker flecks dorsally. The rostral and nasal scales and often the frontonasal are a light yellow or cream. The tail is olive green or brown without other markings. Ventrally yellow-green to light grey-green. The iris is golden yellow to lime green. Natural History Notes: This very large skink (probably the world’s largest) is not often seen because it is nocturnal, sheltering by day in hollows or amongst dense foliage in the larger forest trees; the crevices in strangler fig trees Ficus spp. are favoured habitats. It is almost completely arboreal, rarely encountered on the ground at night as it moves between trees. It is completely herbivorous; while a variety of plants are eaten, it has a marked preference for the young leaves and inflorescence of the aroids of the genus Epipremnum. These climbing epiphytes are common on rainforest trees in the Solomons, and the presence of Corucia can often be determined by the characteristic bite marks on the young leaves. Kinghorn (1928) mentions the anecdotal accounts of Solomon Islanders who claim that Corucia live communally in tree hollows with only the topmost lizard venturing out to feed while the other lizards eat its droppings. This observation is undoubtedly based on the fact that newborn Corucia eat the faeces of the adults in order to establish their culture of the gut flora necessary for the digestion of plant material. During heavy rain after a prolonged dry spell I have seen several Corucia emerge at twilight from hollows in a single fig tree to eagerly lick up the raindrops that had collected on the leaves. Corucia is rather slow moving and generally docile, though when provoked it will elevate its body and exhale with a sharp, loud hiss through its SKINKS, Family SCINCIDAE

49

open mouth, and given the opportunity it will savagely bite its tormentor. It gives birth to a single (rarely two) young that is about a third the size of the adult. Although previously common, despite traditional human predation (Corucia is a prized food item for many rural Solomon Islanders), many populations of Corucia are now threatened by habitat destruction. Also, gross over-collecting of Corucia in the Solomons by avaricious animal dealers supplying the insidious foreign pet trade, has necessitated its CITES (Convention on International Trade in Endangered Species) Appendix II status. Literally thousands of these slow-breeding lizards were exported from the Solomons annually from the mid 1980s until 2002 when trade in the species was stopped following its Appendix II listing. As an example, from the period January 1989 to December 1990 alone, almost 7500 Corucia were exported (Leary, 1991). Understandably, this trade has decimated wild populations and its ongoing effects continue to seriously threaten the longterm survival of many populations of this spectacular skink.

Genus Cryptoblepharus Wiegmann, 1834 Small terrestrial or semi-arboreal skinks. The eye is covered with an immovable transparent scale. Prefrontals well developed. Frontoparietals and interparietal fused into a single shield. Parietals in contact. OCEANIC SNAKE-EYED SKINK Cryptoblepharus poecilopleurus (Wiegmann, 1835) Fig. 28 Distribution: This species occurs widely on many of the islands of the tropical and subtropical Pacific. In the Solomons it has only been recorded from Rennell. Type locality: Pisacoma Island, Peru. Description: Average SVL 45mm, tail about 120% of SVL. Head moderate, snout somewhat elongated and pointed. Limbs well developed and overlapping when adpressed. Supranasals absent, very rarely present. Prefrontals usually in contact, rarely separated. Frontal small, shorter than its distance from the snout, in contact with the first and second of 4 (rarely 3 or 5) supraoculars. The eye is covered with an immovable transparent spectacle. Frontoparietals and interparietal fused into a single shield. Enlarged nuchals present. The two middorsal scale rows are transversely

50

SKINKS, Family SCINCIDAE

enlarged. Rennell populations of this skink have 22 to 28 (usually 24 or 26) midbody scale rows. 18 to 26 lamellae under the fourth toe. Colour: Dorsal ground colour grey-green with a series of heavy darker flecks from the neck to a little past the base of the tail. These dark flecks may coalesce to form irregular longitudinal stripes. An irregular broad, lighter dorsolateral stripe runs from the eye to at least the base of the tail. Laterally dark brown to black with lighter spotting. Head brownish-green with small darker flecks. Tail and limbs grey-green with light and dark spotting. Ventrally white to cream. Natural History Notes: This species occurs on Rennell as scattered populations with high population densities. The lizards live on the bare vertical limestone walls that typify much of Rennell’s coastline (Rennell is an uplifted coral atoll). Where these walls are covered with creeper growth, Cryptoblepharus is replaced by Emoia rennellensis. Cryptoblepharus poecilopleurus feeds almost exclusively on the small moths and isopods that gather in large numbers in crevices in the limestone walls. This species is very active and extremely agile. As soon as the first rays of the morning sun reach the crevices where the lizards shelter nocturnally, they immediately emerge to bask for several minutes until their body temperatures rise sufficiently to allow them to begin their foraging. This lizard displays considerable intraspecific aggression and it is rare to find an individual without a regenerated tail, the original having been lost in tussles with other lizards. Given this species’ wide occurrence further eastwards in Polynesia, and the fact that it does not occur elsewhere in the Solomons, it seems likely that it was introduced to Rennell - as eggs or mature lizards - by the seafaring Polynesians who colonised the island about 1000 years ago.

Genus Emoia Gray, 1845 A large group of small to moderate sized scincid lizards widely distributed in the tropical areas of the Pacific and southeast Asia. Limbs well developed, pentadactyl. 30 or more subdigital lamellae under the fourth toe. Interparietal reduced in size or absent. Supranasals present. Frontoparietals fused. Palpebral disc present in the lower eyelid. Prefrontals separated or narrowly in contact. Parietals in contact. Precloacals usually enlarged. All species are oviparous with most species having a constant clutch size of 2. The majority of Solomon Islands skinks belong to this genus. SKINKS, Family SCINCIDAE

51

KEY TO THE SPECIES OF SOLOMON ISLANDS Emoia 1. Subdigital lamellae smooth and rounded, fewer than 50 beneath the fourth toe: 2 – Subdigital lamellae thin and blade-like, more than 50 beneath the fourth toe: 6 2. A distinct white midlateral stripe from the snout to the groin: jakati – No distinct midlateral stripe: 3 3. Middorsal and/or dorsolateral stripes present, at least on the head and anterior body: caeruleocauda – No middorsal or dorsolateral stripes: 4 4. A small interparietal present: 5 – Interparietal absent, chin and throat yellow: flavigularis 5. Colour grey to grey-green to greenish brown dorsally with a series of wavy whitish transverse bands: atrocostata – Colour uniform black or brown dorsally with or without darker flecks: nigra 6. Middorsal and/or dorsolateral stripes discernible, at least on the head and/or anterior body: 7 – Middorsal and dorsolateral stripes not discernible: 11 7. Midbody scales in 36 or fewer rows: 8 – Midbody scales in 38-42 rows: rufilabialis 8. Distinct middorsal and dorsolateral stripes present from the snout to the tail, tail without dark longitudinal stripes or regular spotting: 9 – Middorsal and/or dorsolateral stripes present on the head and/or part of the body, though sometimes indistinct; if both middorsal and dorsolateral stripes are present for the full length of the body, then the tail with dark longitudinal stripes and/or distinct regular spots, at least anteriorly: 10

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SKINKS, Family SCINCIDAE

9. Parietal eye present (although difficult to discern in some individuals), no fusion of the middorsal pair of scale rows (the middorsal stripe overlaps two adjacent scales for its entire length), tail light brown, grey, blue-grey or greenish: cyanura – Parietal eye absent, some degree of fusion of the middorsal pair of scale rows (the middorsal stripe occupying single scales for at least part of its length), tail blue: impar 10.Middorsal stripe terminating abruptly on the neck, occurring only in the Duff Islands: female taumakoensis – Middorsal stripe not terminating abruptly on the neck, not occurring in the Duff Islands: 14 11. Average adult SVL more than 80mm: 12 – Average adult SVL less than 70mm: 13 12.Midbody scales in 22-28 rows, a dark streak from the snout, through the eye and onto the neck and anterior body, dorsally bronze or greenish bronze: cyanogaster – Midbody scales in 28-34 rows, uniformly bright green with a large black marking on the occiput: sanfordi 13.More than 70 blade-like lamellae beneath the fourth toe, occurring only in the Duff Islands: male taumakoensis – Less than 70 blade-like lamellae beneath the fourth toe, occurring only on Bellona Island: isolata 14.Only occurring on Rennell Island or Makira Island (including Ugi and the Olu Malau group): 15 – Only occurring in the central and western Solomons (from Malaita to Bougainville): 16 15. Midbody scales in 26-32 rows, 65-76 lamellae under the fourth toe, occurring only on Rennell Island: rennellensis – Midbody scales in 28-32 rows, 73-88 lamellae under the fourth toe, occurring only on Makira and neighbouring islands (Olu Malau, Santa Ana, Santa Catalina and Ugi): maculata SKINKS, Family SCINCIDAE

53

16.Midbody scales in 28-34 rows, 73-98 (rarely fewer than 78) lamellae under the fourth toe, occurring only in the central and far western Solomons (Bougainville, Shortland, Choiseul, Isabel, Russell Islands, Guadalcanal, Ngela, Malaita): pseudocyanura – Midbody scales in 30-36 rows, 69-83 lamellae under the fourth toe, occurring only in the central western Solomons (Gizo, Kolombangara, New Georgia, Vangunu): schmidti REEF SKINK Emoia atrocostata (Lesson, 1826) Fig. 29 Brown (1991) refers the Solomon Islands populations to the subspecies freycineti (Duméril and Bibron, 1839) Distribution: Philippines, parts of the Malay Peninsula, Caroline Islands, Indonesia, New Guinea, Cape York in northern Australia, islands of Torres Strait, Solomon Islands. In the Solomons (as the subspecies freycineti): Bougainville, Shortland Islands, Choiseul, Rob Roy, Ranongga, Vella lavella, Gizo, New Georgia, Tetepare, Vangunu, Isabel, Russell Islands, Guadalcanal, Ngela, Malaita, Ontong Java, Makira, Olu Malau Group Islands, Ugi, Rennell, Bellona, Santa Cruz, Reef Islands, Vanikoro. This subspecies also occurs in Vanuatu. Type locality: (for the lectotype Emoia atrocostata freycineti) Vanikoro Island, Solomon Islands. Description: Average SVL 75mm, tail about 150% of SVL. Head moderate, snout narrow and elongate. Limbs well developed, overlapping when adpressed. Body somewhat stout. Prefrontals separated. Supranasals present. Frontal slightly longer than wide, in contact with the first and second of 4 supraoculars. Interparietal distinct. Lower eyelid with a palpebral disc. Enlarged nuchals present. Precloacals slightly enlarged. Midbody scales in 35 to 43 rows. 31 to 42 lamellae beneath the fourth toe. Colour: Dorsally grey to grey-green to almost black with lighter flecks that may tend to form irregular transverse bands. Laterally black with lighter flecks. Ventrally white with a greenish hue. The limbs and tail are marked similarly to the body. Natural History Notes: This is a very common species in the Solomons, occurring only on rocky foreshores bordering the sea where large populations may be present. It is a very active, agile lizard and readily

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SKINKS, Family SCINCIDAE

enters tidal pools to escape potential predators; it can stay submerged in saltwater for several minutes. I have seen these lizards foraging on foreshores where waves are breaking, clinging to the rock while the wave passes over them. Most of their foraging however is done in the intertidal zone where they feed upon small invertebrates such as small crabs, isopods, insects and spiders. Although these lizards will forage on exposed mudflats at low tide, they rarely venture further than about 10 metres from the shelter of the adjacent rocks. Cogger (1992) reports this species as a mangrove inhabitant in northern Australia, but it does not exploit this niche in the Solomons. Its 2 eggs are laid well above the high water line in beach debris, driftwood or hollows in limestone rock. PACIFIC BLUE-TAILED SKINK Emoia caeruleocauda (De Vis, 1892) Figs. 30, 31, 32, 33, 34, 35 This is probably a composite species. Distribution: Marianas, Palau (Caroline Islands), southern Philippines, northern Kalimantan, Maluku Islands (Moluccas), Sulawesi, New Guinea and its island archipelagos, Solomon Islands, Vanuatu and Fiji. In the Solomons: Bougainville, Shortland Islands, Choiseul, Rob Roy, Vella lavella, Ranongga, Gizo, Kolombangara, Rendova, New Georgia, Tetepare, Vangunu, Isabel, Makira, Santa Ana, Santa Catalina, Santa Cruz, Taumako (Duff Islands), Utupua. Brown (1991) also records this species from Bellona and Malaita, though on the basis of my own observations, I believe E. caeruleocauda does not occur on these latter two islands. Type locality for the neotype (Brown, 1991): Rambuso, Sudest Island, Louisiade Archipelago, Papua New Guinea. Description: Average SVL 50mm, tail about 130% SVL. Head moderate, snout short. Limbs well developed, overlapping when adpressed. Supranasals present. Prefrontals separated. A single anterior and posterior loreal. Frontal about 1.5x as long as wide, longer than its distance from the snout, in contact with the first and second of 4 supraoculars. Lower eyelid with a palpebral disc. Frontoparietals and interparietal fused into a single shield. Enlarged nuchals present. Midbody scales in 26 to 36 rows. 30 to 43 smoothly rounded lamellae under the fourth toe. Colour: Dorsally black. A golden middorsal stripe is present, running from the snout to a little posterior to the hindlimbs where it terminates abruptly. SKINKS, Family SCINCIDAE

55

Two golden dorsolateral stripes run from the snout and along the body to the tail where they merge into the bright electric-blue tail colouration. Laterally dark brown to black with lighter flecks. Ventrally grey-white. On Makira and Kolombangara, and also in some New Georgia populations, E. caeruleocauda exhibits marked sexual dichromatism, this is also known in New Guinea populations of E. caeruleocauda. In these dichromatic populations the females are coloured and patterned as above, while the males are golden brown dorsally and the middorsal and dorsolateral stripes may be present on the head and anterior body only, or sometimes indistinct or absent, and the tail is light brown, sometimes with a bluish hue. Natural History Notes: A common terrestrial species. On Makira at least, this species appears to be ecologically intermediate between Emoia cyanura and E. maculata. While E. cyanura almost never enters areas with dense canopy cover, preferring open, cleared areas, E. maculata is a dweller in forests and areas with at least some canopy cover. Emoia caeruleocauda however, utilises both niches equally and in areas such as semi-cleared food gardens or overgrown gardens at the edge of the forest, the three species occur together. In this situation there does not appear to be any overt interspecific interaction apart from the obvious competition for insect food. The presence of one of these species does not preclude the presence of either of the other two. On Bougainville, E. caeruleocauda occurs from the coast up to 1200m elevation (Webster 1969). This lizard is essentially terrestrial though it will forage in low scrub and climb a little distance up the trunks of trees. Its 2 eggs are laid in ground debris. GREEN-BELLIED TREE SKINK Emoia cyanogaster (Lesson, 1826) Figs. 36, 37 Distribution: Bismarck Archipelago (Papua New Guinea), Solomon Islands, Vanuatu. In the Solomons: Bougainville, Shortland Islands, Fauro, Mono, Choiseul, Vella lavella, New Georgia, Tetepare, Vangunu, Isabel, Guadalcanal, Ngela, Malaita, Rennell, Ugi, Olu Malau, Makira, Santa Ana, Santa Cruz, Reef Islands, Utupua, Vanikoro, Tikopia. Type locality: “Oualan” (Kosrae Island), Caroline Islands (in error, Brown, 1991). Description: Average SVL 85mm, tail about 250% of SVL. Head depressed, snout long and slender. Limbs well developed, overlapping

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when adpressed. Supranasals present. Prefrontals in contact or narrowly separated. A single anterior and posterior loreal. Frontal 2x as long as wide, longer than its distance from the snout, in contact with the first and second of 4 supraoculars. Lower eyelid with a palpebral disc. A small interparietal is present. Midbody scales in 22 to 28 rows. 70 to 95 very thin, blade-like lamellae under the fourth toe. Colour: Dorsally golden bronze or greenish bronze with or without lighter and darker flecks. A black streak runs from the nostril, through the eye and onto the neck, continuing in a lateral position until it fades out around midbody. Ventrally yellow-green to bright lime green, usually sharply differentiated from the bronze lateral colouration. Tail marked similarly to the body. Natural History Notes: A very active diurnal lizard. Largely arboreal, it does forage to some extent on the ground. It inhabits forests, domestic gardens and plantations and shows a preference for dwelling in low scrub or on vine and creeper covered trees and tends to avoid larger, baretrunked trees. Its average perch height varies from 1 to 6m. It feeds upon insects and spiders and also smaller lizards. It lays its 2 eggs in ground debris or loose soil or in rotting timber on or above the ground. It is quite common over most of its range in the Solomons. BROWN-TAILED COPPER-STRIPED SKINK Emoia cyanura (Lesson, 1826) Figs. 38, 39 Distribution: Mariana and Caroline Islands, most of tropical Oceania from the Admiralty Islands and Bismarck Archipelago (Papua New Guinea) to Clipperton Island off the west coast of South America. In the Solomons: Bougainville, Shortland Islands, Taro Island in Choiseul Bay (but apparently not mainland Choiseul), Vella lavella, Gizo, Tetepare, Ranongga, Vangunu, Isabel, Russell Islands, Guadalcanal, Ngela, islands of Langalanga Lagoon on western Malaita (but apparently not mainland Malaita), Ontong Java, Makira, Ugi, Olu Malau, Santa Ana, Santa Catalina, Vanikoro, Tikopia, Anuta. Type locality: Tahiti. Description: Average SVL 50mm, tail about 130% of SVL. Head moderate, snout somewhat short. Limbs well developed, overlapping when adpressed. Supranasals present. Prefrontals usually separated, occasionally narrowly in contact. A single anterior and posterior loreal. Frontal 1.5x as long as wide, SKINKS, Family SCINCIDAE

57

longer than its distance from the snout, in contact with first and second of 4 supraoculars. Lower eyelid with a palpebral disc. Frontoparietals and interparietal fused into a single shield. Parietal eye present. Enlarged nuchals present. Precloacals slightly enlarged. Midbody scales in 25 to 32 rows. 58 to 86 (usually around 65 to 70) very thin lamellae under the fourth toe. Colour: Dorsal ground colour grey to brown to almost black. A pale golden middorsal stripe runs from the snout and along the body to the tail where it fades out a little past the level of the hindlimbs. The borders of this stripe are distinct for most of its length. Dorsolateral stripes of equal width and distinction are present from the snout to at least the base of the tail. Laterally grey-brown with lighter and darker flecks. Ventrally ivory white. Tail light brown, grey, blue-grey or greenish. Natural History Notes: This lizard occurs only in dry, marginal areas such as gardens and plantations and along the seashore in areas of creeper growth. It almost never enters forests or areas with closed canopy. It is usually abundant wherever it occurs. It is terrestrial in its habits though it forages over fallen timber and sometimes climbs a little distance up small trees and shrubs. It shows a high tolerance to hot, dry conditions and can be seen basking and foraging where other lizards could not tolerate the high temperatures. It feeds upon small insects and spiders. It lays its 2 eggs in ground debris or hollows in fallen timber and rocks. YELLOW-THROATED SKINK Emoia flavigularis Schmidt, 1932 Fig. 40 Distribution: Endemic to the Solomon Islands: Bougainville, Choiseul, Shortland Island, Fauro, New Georgia, Isabel, Ngela, Malaita. Type locality: Isabel Island, Solomon Islands. Description: Average SVL 70mm, tail about 150% of SVL. Head moderate. Limbs well developed and overlapping when adpressed. Supranasals present. Prefrontals in contact or narrowly separated. A single anterior and posterior loreal. Frontal about 1.5x as long as wide, in contact with the first and second of 4 supraoculars. Lower eyelid with a palpebral disc. Frontoparietals fused into a single shield. Interparietal consistently absent. Enlarged nuchals present. Precloacals slightly enlarged. Midbody scales in 34 to 40 rows. 36 to 48 rounded lamellae under the fourth toe.

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Colour: Golden brown dorsally with darker flecks, particularly on the posterior dorsum and tail where they sometimes form vague transverse bands. Laterally a slightly darker brown with a few yellowish spots on the side of the neck and lighter and darker flecks on the sides of the body. The throat and underside of the neck to the level of the forelimbs are bright yellow. Ventrally grey suffused with yellow. The whole body has a strong metallic gloss. The limbs, particularly the hands and feet, are blackish. Natural History Notes: This species closely resembles Emoia nigra in appearance and habits and can easily be mistaken for it in the field. It is common on Ngela, though apparently uncommon to rare elsewhere in its range. It tends to be an inhabitant of thick, undisturbed forest where it basks in patches of sunlight on the forest floor; it does not appear to occur outside forests or areas with extensive canopy cover. It is a very active, swift-moving lizard and although largely terrestrial in its habits, it will forage some distance up trees and shrubs and will also climb trees to escape predators. BLUE-TAILED COPPER-STRIPED SKINK Emoia impar (Werner, 1898) Fig. 41 Distribution: Bismarck Archipelago (Papua New Guinea), Northern Mariana Islands, Caroline Islands, Marshall Islands, Solomon Islands, Vanuatu, Fiji, the Samoas, Tonga, Cook Islands, Tokelau Islands, Tuamotu Archipelago, Marquesas Islands, Hawai’i. In the Solomons it occurs on Bougainville, Shortland, Mono and Fauro. Type locality (for the neotype designated by Ineich and Zug, 1991): Mioko Island, Bismarck Archipelago, Papua New Guinea. Description: Average SVL 55mm, tail about 180% of SVL. Head moderate. Limbs well developed and overlapping when adpressed. Supranasals present. A single anterior and posterior loreal, the height of the anterior loreal is greater than its length. Parietal eye consistently absent. There is some degree of fusion of the middorsal pair of scale rows. This fusion results in the middorsal stripe occupying a single scale (rather than overlapping two adjacent scales) for at least part of its length. Ineich and Zug (1991) report the neotype as having 29 midbody scale rows and 74 lamellae under the fourth toe of the right foot. SKINKS, Family SCINCIDAE

59

Colour: Dorsal ground colour dark brown to black. Golden middorsal and dorsolateral stripes present and distinct from the snout to at least the level of the hindlimbs. The tail is bright blue in most animals, though very occasionally it may be green or brown. Ventrally a shade of grey. Natural History Notes: This species is very similar to Emoia cyanura, a species with which it is sympatric over a large part of its range in Oceania, and consequently it has long been regarded as a synonym of E. cyanura. However, in addition to distinguishing physical characters (Ineich and Zug, 1991), the two species have been shown to differ in habitat preferences (Bruna et al., 1996). Essentially E. impar frequents forested and shaded areas to a much greater extent than does E. cyanura, and is uncommon in the open, sunny areas favoured by E. cyanura. BELLONA SKINK Emoia isolata Brown, 1991 Fig. 42 Distribution: Endemic to the Solomon Islands: Bellona Island. Type locality: Bellona Island, Solomon Islands. Description: Average SVL 60mm for males, 50mm for females, tail about 120% of SVL. Head moderate, somewhat depressed, snout slender. Limbs well developed, overlapping when adpressed. Supranasals present, in contact with the anterior loreal. Prefrontals narrowly separated. Lower eyelid with transparent palpebral disk. Midbody scales in 31 to 34 rows. 56 to 64 blade-like lamellae under the fourth toe. Colour: Dorsal ground colour light to medium brown with a sparse scattering of darker flecks. Laterally darker brown with light flecks, this lateral colour fairly sharply differentiated from the lighter dorsum. On some specimens vague, light dorsolateral lines run from the snout to a little anterior to the hind limbs. A vague middorsal light stripe may also be present on the head, running from the snout and fading out before it reaches the neck. Ventrally light yellowish cream to light orange, becoming slightly darker in the region of the vent. An ill-defined dark stripe runs from the snout through the eye and terminates a little anterior to the ear opening. Tail uniformly medium brown above becoming lighter on the sides and light yellowish grey below. Limbs dark brown with sparse lighter flecks.

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Natural History Notes: Very common on Bellona in both forested and semi-cleared areas with some canopy cover. This lizard is a very active, diurnal animal and feeds on a variety of small arthropods. It is essentially terrestrial though it does forage in low shrubs and on the trunks of larger trees. Two eggs are laid in any suitable cover, such as in decaying humus or rotting timber on the forest floor. Emoia jakati (Kopstein, 1926) Fig. 43 Distribution: Caroline Islands, Marshall Islands, New Guinea and adjacent island archipelagos, Solomon Islands. In the Solomons it occurs only on Bougainville. Type locality (for the lectotype [Brown, 1991]): Jakati River region, Vogelkop Peninsula, Irian Jaya, Indonesia. Description: Average SVL 45mm for males, 50mm for females, tail about 130% SVL. Head moderate, snout short and bluntly rounded. Limbs well developed, overlapping when adpressed. Supranasals present. Prefrontals separated. A single anterior and posterior loreal. Frontal about 1.5x as long as wide, longer than its distance from the snout, usually in contact with the first and second (rarely the third) of 4 supraoculars. Lower eyelid with a palpebral disc. Frontoparietals and interparietal fused into a single shield. A single pair of enlarged nuchals present. Midbody scales in 29 to 39 rows, 30 to 42 smoothly rounded lamellae under the fourth toe. Colour: Dorsally light to medium brown. Light middorsal and dorsolateral stripes of varying width and distinction and edged with black are usually present, though sometimes absent. Laterally dark brown to almost black. A white midlateral stripe is present from the snout to the groin. An ill-defined lighter ventrolateral stripe is occasionally present. The tail is marked similarly to the body. Natural History Notes: A small terrestrial species that is common in both forested and cultivated areas. It also occurs in suburban gardens in many towns in Papua New Guinea. The species is reported as occurring from sea level to 1800m in parts of Papua New Guinea (Brown, 1991). Like most terrestrial Emoia, E. jakati lays its 2 eggs in any suitable ground debris. On Bougainville this species is common in coconut plantations, laying its eggs amongst the piles of discarded husks.

SKINKS, Family SCINCIDAE

61

SPOTTED BLUE-TAILED SKINK Emoia maculata Brown, 1953 Figs. 44, 45 Distribution: Endemic to the eastern Solomon Islands of Makira, Olu Malau, Santa Ana, Santa Catalina and Ugi. Type locality: Ugi Island, Solomon Islands. Description: Average SVL 55mm, tail about 150% of SVL. Head moderate. Body a little slender. Limbs well developed and overlapping when adpressed. Supranasals present. Prefrontals separated or narrowly in contact. A single anterior and posterior loreal. Frontal 1.5x as long as wide, longer than its distance from the snout, in contact with the first and second of 4 supraoculars. Lower eyelid with a palpebral disc. Frontoparietals and interparietal fused into a single shield. Enlarged nuchals present. Precloacals slightly enlarged. Midbody scales in 28 to 32 rows. 73 to 88 very thin, blade-like lamellae under the fourth toe. Colour: Dorsal ground colour brown to black, often with a bluish or greenish hue posteriorly. A light middorsal stripe may be present from the snout to around the base of the tail. This stripe is diffuse or absent in some animals. A broad, light dorsolateral stripe runs from the snout, above the eye and along the body for most of its length, though the borders of this stripe are usually diffuse posteriorly. Below the dorsolateral stripe a broad black stripe runs from the snout, through the eye, and along the body in a midlateral position to the groin. The sides of the neck and the throat are orange. Ventrally yellow to dull orange. The tail is a dull to bright blue and usually marked with large black flecks arranged into regular pairs. Natural History Notes: Emoia maculata is essentially an inhabitant of lowland forest and does not usually enter areas without some canopy cover, though it is sometimes encountered foraging around the edges of cultivated areas. It is a semi-arboreal species, foraging over leaf litter on the forest floor and climbing up to 2 or 3m on small trees and creepers in search of its insect food. It is a very active, agile species; I once observed an individual on Malaupaina Island in the Olu Malau group drop from a branch more than 2m high to grab an insect that it had observed moving on the ground. This species is abundant throughout its range and in some areas high population densities occur. It lays its 2 eggs in ground debris or in rotting fallen timber.

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PACIFIC BLACK SKINK Emoia nigra (Jacquinot and Guichenot, 1853) Figs. 46, 47, 48 Distribution: Bismarck Archipelago (Papua New Guinea), Solomon Islands, Vanuatu, Fiji, Samoa, Tonga. This is the widest ranging reptile in the Solomons, occurring on virtually every island of the group from Bougainville to the far Eastern Outer Islands of Tikopia and Anuta. Type locality not recorded, holotype lost. Description: Average SVL 100mm, tail about 150% of SVL. Head and body moderately large. Body somewhat stout. Limbs well developed and overlapping when adpressed. Supranasals present. Prefrontals in contact or narrowly separated. A single anterior and posterior loreal. Frontal about 2x as long as wide, longer than its distance from the snout, in contact with first and second of 4 supraoculars. Lower eyelid with a palpebral disc. Frontoparietals fused into a single shield. A small interparietal is present. Enlarged nuchals present. Precloacals slightly enlarged. Midbody scales in 33 to 40 rows. 32 to 39 rounded lamellae under the fourth toe. Colour: Uniformly glossy black or brown above, sometimes with a scattering of lighter flecks laterally. Ventrally whitish to grey or yellowish to dull orange. Juveniles are dark grey or brown with indistinct darker flecks dorsally and distinct whitish spots laterally. On Guadalcanal E. nigra are light brown to dark brown dorsally with darker flecks and darker brown to almost black laterally with lighter flecks. Ventrally they are white to dull cream. Juveniles from Guadalcanal are a light golden dorsally, with or without darker flecks and dark brown laterally with light spots. Natural History Notes: A very active, robust diurnal lizard. It is common in the Solomons and occurs with some abundance in many areas. Although it forages mainly on the ground, it is at least partly arboreal and will often take refuge up the nearest tree if alarmed on the ground. It occupies a wide variety of habitats from forests to domestic gardens and plantations and along the seashore in areas with creeper growth. It feeds upon insects and spiders and smaller lizards. A large resident population of E. nigra lived in the gardens around my house in Honiara on Guadalcanal, and on several occasions I saw them pursue, catch and eat E. pseudocyanura. E. nigra lays from 2 to 4 eggs in a clutch, usually in any suitable ground debris, the bases of arboreal ferns, or debris collected in tree forks. SKINKS, Family SCINCIDAE

63

SOLOMONS BLUE-TAILED SKINK Emoia pseudocyanura Brown, 1991 Figs. 49, 50, 51, 52, 53 This is probably a composite species. Distribution: Endemic to the Solomon Islands: Bougainville, Shortland, Choiseul, Isabel, Russell Islands, Guadalcanal, Ngela, Malaita. Type locality: Honiara, Guadalcanal Island, Solomon Islands. Description: Average SVL 55mm, tail about 160% of SVL. Malaita populations of E. pseudocyanura (which may be a undescribed species) show sexual dimorphism in SVL, the males being larger than the females and growing up to 70mm SVL. Head moderate, snout with a fairly sharp taper, rounded at its tip. Limbs well developed, overlapping when adpressed. Supranasals present. Prefrontals separated. A single anterior and posterior loreal. Frontal about 2x as long as wide, longer than its distance from the snout, in contact with the first and second of 4 supraoculars. Lower eyelid with a palpebral disc. Frontoparietals and interparietal fused into a single shield. Enlarged nuchals present. Midbody scales in 28 to 34 rows. 73 to 96 thin, blade-like lamellae under the fourth toe. Colour: The colour and pattern of E. pseudocyanura is quite variable. Dorsal ground colour very dark brown to black. In most populations a middorsal stripe runs from the snout to around the base of the tail. In some populations, most noticeably those from Malaita, the middorsal stripe may be distinct only on the head and neck, usually fading out before midbody (fig. 53). In populations on Shortland Island, the middorsal stripe often continues unbroken well onto the tail (fig. 49). Pale dorsolateral stripes run from the snout to the base of the tail, usually widening and becoming more diffuse posteriorly. Laterally with a broad, black longitudinal band with lighter flecks. Shortland Island specimens often have a narrow yellow midlateral line running from the neck to the groin (fig. 49). The tail is dull to bright blue or blue-green, usually with a regular series of paired black spots for most of its length. The spotting on the tail is absent in Malaita populations. Ventrally creamy white to dull yellow. Natural History Notes: Emoia pseudocyanura is a semi-aboreal lizard that occurs abundantly over most of its range. While it does inhabit forested areas from sea level up to around 1000m elevation on the larger islands, it is essentially a forest-edge species and often frequents cultivated

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areas and domestic gardens, provided there is ample tree cover. It is a highly active species and in areas where large population densities occur, there is frequent intraspecific aggression relating to established territories with interlopers being chased for considerable distances by resident lizards. Females lay 2 eggs in ground debris such as in and under rotting fallen timber. RENNELL BLUE-TAILED SKINK Emoia rennellensis Brown, 1991 Fig. 54 Distribution: Endemic to the Solomon Islands: Rennell Island. Type locality: Rennell Island, Solomon Islands. Description: Average SVL 45mm, tail about 130% of SVL. Head moderate, snout short. Limbs well developed, overlapping when adpressed. Supranasals present. Prefrontals separated. A single anterior and posterior loreal. Frontal about 1.5x as long as wide, longer than its distance from the snout, in contact with the first and second of 4 supraoculars. Lower eyelid with a palpebral disc. Frontoparietals and interparietal fused into a single shield. Enlarged nuchals present. Midbody scales in 26 to 32 rows. 65 to 76 thin, blade-like lamellae under the fourth toe. Colour: Very similar in colour and pattern to Emoia pseudocyanura. The dorsal ground colour is dark brown to black, a lighter brown on the head. An ill-defined, diffuse, lighter stripe is usually present middorsally. Golden dorsolateral stripes are present from the snout to least midbody. A broad black stripe runs from the snout, through the eye, and along the body in a midlateral position to the groin. A regular scattering of large light spots are present in this dark band. Ventrally yellowish-white to cream. The tail is dull to bright blue with a series of paired black spots along most of its length. Natural History Notes: A common skink on Rennell, where it occurs in closed canopy forest and cleared and cultivated areas. It also lives amongst the creeper growth on the vertical limestone cliffs that make up most of Rennell’s coast. It is an active, agile lizard and partly arboreal in its habits.It lays its 2 eggs in ground debris and also in detritus-filled limestone crevices. SKINKS, Family SCINCIDAE

65

RED-LIPPED SKINK Emoia rufilabialis McCoy and Webber, 1984 Figs. 55, 56 Distribution: Endemic to the Solomon Islands: Santa Cruz Island. Type locality: Graciosa Bay, Santa Cruz Island, Solomon Islands. Description: Average SVL 60mm, tail about 150% of SVL. Head moderate, snout sharply tapering. Limbs well developed, overlapping when adpressed. Supranasals present. Prefrontals separated. A single anterior and posterior loreal. Frontal about 1.5x as long as wide, longer than its distance from the snout, in contact with the first and second of 4 supraoculars. Lower eyelid with a palpebral disc. Frontoparietals and interparietal fused into a single shield. Enlarged nuchals present. Midbody scales in 36 to 42 rows. 68-84 thin, blade-like lamellae under the fourth toe. Colour: Dorsal ground colour dark brown to black. The snout and lips are dull to bright orange-red. A light cream middorsal stripe may be present from the snout to the base of the tail. This stripe may be absent in old males but is usually present, at least on the head. Light cream dorsolateral stripes may be present or absent as per the middorsal stripe. The striping is much more pronounced in females and juvenile males. Laterally yellow to greenish-gold with an irregular peppering of dark spots. Ventrally dull yellow to light orange. Tail dull blue to grey-green with darker spotting, this spotting often arranged into distinctive pairs in females and juvenile males. Natural History Notes: A common species on Santa Cruz. It occurs most abundantly in areas with some canopy cover, such as overgrown food gardens and coconut plantations. It does not occur in open areas lacking overhead cover, nor is it commonly found in closed canopy forest. It is an active, agile lizard, and although largely terrestrial, it does forage and perch up to 2m above the ground. SANFORD’S TREE SKINK Emoia sanfordi Schmidt and Burt, 1930 Fig. 57 Distribution: Vanuatu, Solomon Islands. There is no record of this lizard having been collected in the Solomons since it was originally described in 1930 (Schmidt and Burt, 1930) when two specimens from Fauro Island in

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SKINKS, Family SCINCIDAE

the far western Solomons were noted as paratypes. Furthermore the Fauro people do not recognise the animal from colour photographs and it is probable that the reported provenance of the Fauro paratypes is in error. Type locality: Elephant and Espiritu Santo Islands, Vanuatu. Description: Average SVL 100mm, tail about 180% of SVL. Head slightly depressed, snout somewhat slender. Body stout. Limbs well developed and overlapping when adpressed, digits very strongly clawed. Supranasals present. Prefrontals in contact. A single anterior and posterior loreal. Frontal longer than wide, a little longer than its distance from snout, in contact with first and second of 4 supraoculars. Lower eyelid with a palpebral disc. Frontoparietals fused into a single large shield. A small interparietal is present. Enlarged nuchals present. Midbody scales in 28 to 34 rows. 61 to 76 thin, blade-like lamellae under the fourth toe. Colour: Uniformly bright green with a distinctive, irregularly-shaped black blotch in the occipital region and a very sparse scattering of black flecks on the body and tail. Ventrally greenish-white. The iris of the eye is green with golden flecks. Natural History Notes: I have observed wild populations of this large and spectacular arboreal Emoia on Efate Island in Vanuatu. It is a very distinctive skink due to its size, its highly active habits and its brilliant green colouration. It is an abundant species, particularly in semi-cleared rural areas and in overgrown food gardens. It also occurs in closed canopy forest. Brown (1991) mentions gravid female E. sanfordi as having from 3 to 5 oviducal eggs. SCHMIDT’S SKINK Emoia schmidti Brown, 1953 Figs. 58, 59 Distribution: Endemic to the western Solomon Islands of Gizo, Kolombangara, New Georgia, Tetepare, Vangunu and nearby smaller islands. Type locality: New Georgia Island, Solomon Islands. Description: Average SVL 55mm, tail about 150% of SVL. Head moderate, snout tapered and bluntly rounded. Limbs well developed, overlapping when adpressed. Supranasals present. Prefrontals separated. A single anterior and posterior loreal. Frontal about 2x as long as wide, longer than its distance from the snout, in contact with the first and second of 4 supraoculars. Lower eyelid SKINKS, Family SCINCIDAE

67

with a palpebral disc. Frontoparietals and interparietal fused into a single shield. Enlarged nuchals present. Midbody scales in 30 to 36 rows. 69 to 83 thin, blade-like lamellae under the fourth toe. Colour: Dorsally dark brown to almost black, lighter on the head, particularly the snout. No middorsal stripe. An ill-defined dorsolateral stripe is usually present from the snout to around midbody where it diffuses into the ground colour. Laterally very dark brown to black with a scattering of lighter flecks. The tail is grey to blue-green, usually with darker markings which in some populations may be arranged into paired spots. Ventrally cream to dull yellow. Natural History Notes: Emoia schmidti is a very common lizard in both closed canopy forests and cultivated tracts with some tree cover. It does not occur in open cleared areas. It is partly arboreal with a perch height up to about 2m, though most of its foraging is done on the ground where it searches for its arthropod prey on and under the leaf litter. Females lay their 2 eggs in suitable ground debris in shaded areas. TAUMAKO SKINK Emoia taumakoensis McCoy and Webber, 1984 Figs. 60, 61 Distribution: Endemic to the Solomon Islands: Duff Islands. Type locality: Taumako, Duff Islands, Solomon Islands. Description: Average SVL 55mm, tail about 150% SVL. Head moderate, snout somewhat short. Body a little robust. Limbs well developed, overlapping when adpressed. Supranasals present. Prefrontals separated. A single anterior and posterior loreal. Frontal about 2x as long as wide, longer than its distance from the snout, in contact with the first and second of 4 supraoculars. Lower eyelid with a palpebral disc. Frontoparietals and interparietal fused into a single shield. Enlarged nuchals present. Midbody scales in 32 to 35 rows. 76 to 85 thin, blade-like lamellae under the fourth toe. Colour: This is a sexually dichromatic species. Males: dorsally light to dark olive brown, a little darker on the head. The side of the neck is light pink. The body is uniform in colour, there are no longitudinal stripes. The tail is a uniform blue-green. Females: Dorsal ground colour dark brown. A golden middorsal stripe runs from the snout to the neck where it terminates abruptly. Golden dorsolateral stripes run from the snout to the

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base of the tail, their margins diffuse posteriorly. Tail light blue to bluegreen. Ventrally both males and females are cream to dull yellow. Natural History Notes: An inhabitant of semi-closed canopy forest and partially cleared areas. It is an active skink and forages for its insect food through leaf litter in forested areas and also amongst the heaped up vegetable scraps at the boundaries of local villages. When basking it tends to utilise high spots in its environment - such as shrubs, boulders and the buttress roots of large trees.

Genus Eugongylus Moderate sized terrestrial lizards. Lower eyelid scaly. Supranasals present. Prefrontals well developed, usually separated. Frontoparietals and interparietal distinct. Parietals in contact. Limbs moderate, just meeting or just failing to meet when adpressed. KEY TO THE SOLOMON ISLANDS SPECIES OF Eugongylus – Midbody scales in 34-38 rows, not occurring on Rennell Island: albofasciolatus – Midbody scales in 28-30 rows, only occurring on Rennell Island: rufescens Eugongylus albofasciolatus (Günther, 1872) Figs. 62, 63 Distribution: New Guinea and the Bismarck Archipelago, Solomon Islands. In the Solomons: Bougainville, Shortland Islands, Fauro, Choiseul, Vella Lavella, New Georgia, Vangunu, Isabel, Guadalcanal, Ngela, Malaita, Makira, Ugi, Olu Malau, Santa Ana, Santa Cruz, Taumako (Duff Islands). Burt and Burt (1932) recorded this species from Tikopia, though the Tikopian people, on being shown a photograph of this lizard, informed me that it does not occur on their island. Description: Average SVL 160mm, tail about 140% of SVL. Head moderate, snout short. Body somewhat elongated and stout. Limbs rather short, just meeting when adpressed. The base of the tail is almost as thick as the body. Supranasals present. Prefrontals separated. A single anterior and posterior loreal. Frontal 2x as long as wide, longer than its distance from the SKINKS, Family SCINCIDAE

69

snout, in contact with first and second of 4 (sometimes 5) supraoculars. Lower eyelid scaly. Frontoparietals and interparietal distinct. External ear opening with several small overlapping lobules anteriorly. Midbody scales 34 to 38 rows. 16 to 22 lamellae under the fourth toe. Colour: Dorsally light to dark brown with lighter transverse bands that are most distinct on the anterior half of the body. The lighter banding is more pronounced in juveniles. The lips are barred with dark streaks which continue onto the throat. Ventrally cream without other markings. The whole body has a strong iridescent sheen. The tail is uniformly a shade of brown, though sometimes with indistinct lighter banding at its base . Natural History Notes: A reasonably common species. It is occasionally found moving about during the day during overcast conditions or in areas of heavy shade, otherwise it tends to a crepuscular or nocturnal existence. Mainly terrestrial, though I have seen it foraging on the buttress roots of large forest trees at night. It shelters in leaf litter and under fallen timber on the forest floor and in crevices among the root systems of large trees during the day. It is sometimes found under piles of coconut husks in plantations. I have also seen E. albofasciolatus take refuge down the burrows of land crabs. Food consists of insects and spiders, small land crabs and smaller lizards. It is oviparous, laying up to 5 eggs. Eugongylus rufescens (Shaw, 1802) Fig. 64 Distribution: Maluku (Moluccas), New Guinea, Admiralty Islands (Papua New Guinea), northern Cape York Peninsula and the Torres Strait Islands in Australia, Solomon Islands. In the Solomons it has only been found on Rennell Island. Description: Average SVL 120mm, tail about 120% of SVL. Head moderate. Body elongated and stout. Limbs short, not meeting when adpressed. Supranasals present. Prefrontals usually separated. A single anterior and posterior loreal. Frontal about 2x as long as wide, in contact with the first and second of 4 supraoculars. Lower eyelid scaly. Frontoparietals and interparietal distinct. Ear opening with several small overlapping lobules anteriorly. Midbody scales in 28 to 30 rows. In young specimens the scales are sometimes multicarinate. 16 to 21 lamellae under the fourth toe.

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Colour: Adults uniformly light to dark brown above with a marked iridescent sheen. Lips barred with faint dark bars which many continue onto the throat. Ventrally yellowish. Juveniles are maroon to red-brown with a series of narrow white transverse bands. Natural History Notes: This species is quite common on Rennell. Its habits are similar to those of E. albofasciolatus excepting that it tends to be more active diurnally, and occasionally forages in areas of open sunlight. It shelters under fallen timber and limestone rock in the forest on Rennell. Food probably consists of small arthropods and small lizards.

Genus Geomyersia Greer and Parker, 1968 Body depressed. Prefrontals absent. A single median internuchal scale present between a pair of nuchals. Lower eyelid with a palpebral disc. Geomyersia glabra Greer and Parker, 1968 Fig. 65 Distribution: Endemic to the Solomon Islands: Bougainville, Ngela. Type locality: Mutahi, Bougainville, Papua New Guinea. Description: Average SVL 35mm, tail about 100% of SVL. Head very small, snout short. Body depressed, elongated. Limbs short, failing to meet when adpressed. Digits pentadactyl. Prefrontals absent. A single loreal is present. Frontal broadly in contact with the frontonasal and the first and second of 4 supraoculars. Lower eyelid with a palpebral disc. Frontoparietals fused into a single shield. Interparietal distinct. A single pair of nuchals present, separated by a small median scale. Median precloacals enlarged. Midbody scales in 22 to 24 rows. 11 to 15 lamellae under the fourth toe. Colour: Body and tail very glossy with a pronounced iridescent sheen. Dorsally a shade of light or dark brown. A series of anteriorly directed dark chevron markings are present from the neck and along the body in a middorsal position and along the top of the tail. A broken dark line runs from behind the eye and along the body in a dorsolateral position and onto the tail. Laterally dark brown to black. Ventrally grey with darker spots which become thicker on the posterior venter and underside of the tail. The iris is bright orange. There are usually light cream sutures between the supralabials and the parietals are often lighter in colour than the head and dorsum. SKINKS, Family SCINCIDAE

71

Natural History Notes: Observations I have made of both wild and captive specimens of this small lizard indicate that it is nocturnal and a burrower which no doubt accounts in part for its apparent rarity. Specimens I have found on Ngela have been sheltering under fallen timber and stones in moist conditions in river valleys in rainforest at about 220m elevation. On Bougainville it has been found from 150m to over 1000m. It is oviparous and apparently lays a single egg at a time (Greer and Parker 1968b). Food presumably consists of very small arthropods.

Genus Lamprolepis Fitzinger, 1843 Small to moderate sized skinks. Lower eyelid scaly. Limbs well developed, overlapping when adpressed. Frontoparietals and interparietal distinct. Enlarged nuchal scales present. Parietals in contact. EMERALD TREE SKINK Lamprolepis smaragdina (Lesson, 1830) Figs. 66, 67 Distribution: Philippines, Sulawesi, Maluku (Moluccas), the Lesser Sunda Islands west of Bali, Caroline Islands, Marshall Islands, New Guinea and adjacent island archipelagos, Solomon Islands. In the Solomons: Bougainville, Shortland, Fauro, Mono, Choiseul, Ranongga, Vella lavella, Gizo, Kolombangara, New Georgia, Tetepare, Vangunu, Isabel, Russell Islands, Malaita, Ontong Java, Santa Cruz, Reef Islands, Taumako (Duff Islands), Vanikoro, Utupua, Tikopia. It also occurs as introduced breeding populations in and around the botanical gardens area in Honiara on Guadalcanal. Type locality: Caroline Islands. Description: Average SVL 100mm for males, 85mm for females, tail about 120% of SVL. Head moderate, somewhat depressed, snout slender. Limbs well developed, overlapping when adpressed. Supranasals absent. Prefrontals usually separated, sometimes in contact. A single anterior and posterior loreal. Frontal 1.5x as long as wide, longer than its distance from the snout, in contact with the first and second of 4 supraoculars. Lower eyelid scaly. Frontoparietals and interparietal distinct. Ear opening with 2 or 3 small overlapping lobules anteriorly. Enlarged nuchals present. Precloacals enlarged. A distinctive enlarged scale is present on the heel of the

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hindlimb, this scale is more pronounced in males. Midbody scales in 22 to 24 rows. 26 to 35 lamellae under the fourth toe, the distal lamellae compressed. Colour: Bright green dorsally and laterally, usually with darker flecks. In some populations the posterior body and tail is a light golden-green. Ventrally yellow to yellow-green to greenish white. In most populations the base of the tail around the point of insertion of the hind limbs and the hind limbs themselves are a light olive green or brown with large light spots and smaller dark flecks. Lizards from Ontong Java are drab olive dorsally with lighter and darker flecking, the ground colour lighter on the head and anterior body. This olive colouration is stated by Greer (1970) to be indicative of more primitive populations. Natural History Notes: This distinctive diurnal skink is almost totally arboreal although it comes to the ground infrequently to forage. It is very swift moving and highly agile. It feeds upon a variety of arthropods and also flowers - both petals and nectar - and fruits. Its preferred habitat is both large and small bare-trunked trees in semi-cleared areas, and also in domestic gardens. It is almost never found in closed-canopy forest. It is also a common inhabitant of the trunks and inflorescence of coconut palms in some areas. Normal resting/basking position on tree trunks is typically head down with neck and head extended out to a near-horizontal position. The head is pressed against the substrate and the animal moves around the trunk when a potential threat is perceived. Males are highly territorial and will vigorously attack any would-be interloper. Oviparous, its 2 eggs are laid in rotting timber or other detritus on the ground or in the humus collected in tree forks. It is quite common over most of its range in the Solomons (though uncommon on Malaita) and particularly abundant in some areas such as Santa Cruz and the Reef Islands.

Genus Lipinia Gray, 1845 Small, slender lizards, usually marked with distinctive longitudinal striping. Head depressed. Supranasals present. Lower eyelid with a palpebral disc. Frontoparietals distinct or fused. Median precloacals enlarged.

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73

MOTH SKINK Lipinia noctua (Lesson, 1830) Fig. 68 Distribution: Caroline Islands, Marshall Islands, Maluku (Moluccas), New Guinea and nearby islands, Solomon Islands, Vanuatu, most of tropical Oceania east to Hawai’i and Pitcairn Island. In the Solomons: Bougainville, Shortland Islands, New Georgia, Vangunu, Isabel, Guadalcanal, Ngela, Makira, Ugi, Olu Malau. Type locality: Kosrae, Caroline Islands. Description: Average SVL 45mm, tail about 120% of SVL. Head elongated, depressed, snout rather long and pointed. Limbs well developed, overlapping when adpressed. Supranasals absent. Prefrontals in contact or narrowly separated. A single anterior and posterior loreal. Frontal long and kite-shaped, much longer than its distance from the snout, in contact with the first and second and occasionally the third of 4 supraoculars. Lower eyelid with a palpebral disc. Enlarged nuchals present. Median precloacals enlarged. Midbody scales in 23 to 30 rows. 17 to 25 lamellae under the fourth toe. Colour: Head medium to dark brown with black flecks. A more or less well defined yellow spot is present on the occiput. Continuous with this spot is a yellow middorsal stripe which terminates around the level of the hind limbs. Dorsally golden brown to black. The lips are usually barred with black and white. A pair of yellowish broken dorsolateral lines are usually present from the eye to the level of the hindlimbs. A thick black line runs from the snout, through the eye and along the body in a lateral position, breaking up into a series of heavy spots a little anterior to the hindlimbs. The tail is light brown or orange-brown with lighter and darker flecks. Ventrally white to yellowish. Natural History Notes: This small skink is uncommon in the Solomons. It is at least partly terrestrial; I have found specimens on the ground, including one on Ugi that was sheltering under a log in a cleared area adjacent to a village and one on Malaupaina Island that was sheltering in driftwood at the edge of a rocky beach. On Ngela I have seen L. noctua living in large trees, particularly mango trees, and in Honiara on Guadalcanal, over a period of 5 years, I observed a breeding colony of these skinks living in a fig tree in my backyard. The lizards in this colony, which numbered around 4 to 5 adults and up to 7 or 8 juveniles at any one time, foraged on the bare trunk

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of the tree up to around 3 or 4 metres above the ground and sheltered in the leaf litter that collected in forks on the tree as well as in the trailing dead leaves of commensal arboreal ferns. While they would occasionally forage in sunlit parts of the trunk, they did not appear to bask at all, and the majority of their diurnal foraging was done in the shaded side of the tree. They were also active very early in the morning before the sun had reached the tree. Their food consists of any arthropods small enough to be captured and swallowed, including ants. Lipinia noctua gives birth to 1 or 2 young.

Genus Prasinohaema Greer, 1974 Small to medium sized arboreal skinks. Blood plasma and other tissues dark green. Tail prehensile with glandular scales on its ventral tip. GREEN-BLOODED SKINK Prasinohaema virens (Boulenger, 1883) Figs. 69, 70 Distribution: New Guinea, Solomon Islands. In the Solomons: Bougainville, Shortland Islands, Mono, Choiseul, Vella lavella, New Georgia, Tetepare, Vangunu, Isabel, Guadalcanal, Ngela, Malaita, Makira, Ugi, Olu Malau Islands, Santa Ana, Santa Cruz, Reef Islands, Vanikoro, Taumako (Duff Islands), Utupua, Tikopia. Description: Average SVL 50mm, tail about 120% of SVL. Head elongate, quite depressed. Body slender. Limbs well developed, overlapping when adpressed. Tail prehensile. Supranasals absent. Prefrontals well developed, in contact. A single anterior and posterior loreal. Frontal 2x as long as wide, longer than its distance from snout, in contact with first and second of 5 supraoculars. Lower eyelid with a palpebral disc. A large interparietal, much larger than each frontoparietal. Parietals narrowly in contact behind the interparietal. External ear opening with 3 or 4 lobules at its anterior edge. Midbody scales in 32 to 38 rows. Digits expanded proximally with 14 to 18 lamellae on the expanded portion and 6 to 7 on the compressed portion of the fourth toe. Colour: Uniformly pale bright green to light olive green above. An indistinct vertebral stripe is present in some individuals. Ventrally bright yellow to yellow-green. SKINKS, Family SCINCIDAE

75

Natural History Notes: This small skink is totally arboreal; even if forced to descend down the trunk of a tree to within a short distance of the ground, it will invariably try to escape up the tree rather than seek shelter on the ground. It lives on larger and smaller trees and also woody vines and creepers. Its perch height varies from 2 to 20m or more. It is a forest dweller and rarely enters cleared areas. It is quite active though not as agile as some of the arboreal Emoia for example. When moving, it keeps its head and body pressed close to the substrate. At night it sleeps in an exposed position on a branch or leaf. Its 2 eggs are laid in the decaying leaf litter which collects in tree forks or in the bases of arboreal ferns or in moist humus at the bases of trees. This species and four other members of its genus from New Guinea are unusual among vertebrates in that they possess a green blood pigment. This pigment is thought to be the bile pigment biliverdin or a close chemical relative (Greer and Raizes 1969). This green colouration is apparent when the lizard opens its mouth and the green tongue and mucosa can be seen. The eggs are also green. The subdigital lamellae of this lizard function similarly to those of many gekkonids and enable it to move on smooth vertical surfaces or on the underside of branches, I have seen captive Prasinohaema climb up the wall of a glass aquarium.

Genus Sphenomorphus Fitzinger, 1843 Small to moderate sized terrestrial skinks. Lower eyelid scaly. Frontoparietals well developed. Parietals in contact. Limbs poorly to moderately well developed, pentadactyl. The systematics of this group of skinks in the New Guinea-Solomons region is quite complex and is currently subject to ongoing research. This research will almost certainly result in new species designations for a number of populations in the Solomons. KEY TO THE SPECIES OF SOLOMON ISLANDS Sphenomorphus 1. Head shields partially fragmented: 2 – Head shields not fragmented: 3 2. One to 2 pairs of nuchals, midbody scales in 53-60 rows. Large size (up to 160mm SVL): taylori – No enlarged paired nuchals, midbody scales in 42-46 rows. Moderate size (up to 72mm SVL): fragosus

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3. Enlarged paired nuchal scales present: 4 – Enlarged paired nuchal scales absent: 7 4. Prefrontals in contact, or if separated, then adpressed limbs meet or overlap: 5 – Prefrontals separated, adpressed limbs fail to meet, midbody scales in 22-26 rows, 15-17 lamellae under the fourth toe: solomonis 5. 26 or more scale rows at midbody: 6 – 22-25 scale rows at midbody, 18-21 lamellae under the fourth toe. small size (average SVL 35mm): bignelli 6. Midbody scales in 32-41 rows, 20-29 lamellae under the fourth toe, prefrontals usually in contact, rarely separated, found only in the central and western Solomons: cranei – Midbody scales in 26-30 rows, 18-23 lamellae under the fourth toe, prefrontals in contact, found only in the eastern Solomons: Sphenomorphus sp. 7. Midbody scales in 33 or more rows. Adpressed limbs meet or overlap: 8 – Midbody scales in 28-32 rows. 13-20 lamellae under the fourth toe. Prefrontals in contact. Adpressed limbs do not meet: tanneri 8. Average SVL 120mm, adpressed limbs barely meet: woodfordi – Average SVL 70mm, adpressed limbs overlap: 9 9. Prefrontals in contact, 36-47 midbody scale rows, normally 4 supraoculars: concinnatus – Prefrontals narrowly separated, 36 midbody scale rows, 5 supraoculars (in the unique specimen): transversus

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“Sphenomorphus” bignelli Schmidt 1932 Figs. 71, 72 Although this species was originally described as a Sphenomorphus, and this assignation is maintained here for convenience, it is now known to be incorrect and bignelli will in the future be placed in a different and probably new genus (Glenn Shea, pers. comm.). Distribution: Endemic to Solomon Islands: Kolombangara, New Georgia, Tetepare, Vangunu, Russell Islands, Guadalcanal, Ngela, Malaita. Type locality: Kolombangara, Solomon Islands. A specimen of Sphenomorphus minutus from the Russell Islands (Burt and Burt, 1932) probably refers to this species. Description: Average SVL 35mm, tail about 110% of SVL. Head small, snout pointed. Body slightly elongated. Limbs moderate, just meeting when adpressed. Supranasals absent. Prefrontals well developed, usually separated, sometimes narrowly in contact. A single anterior and posterior loreal. Frontal about 2x as long as wide, in contact with the first and second of 4 supraoculars. Lower eyelid scaly. Interparietal distinct. Enlarged nuchals present. Precloacals slightly enlarged. Midbody scales in 22 to 25 rows. 18 to 21 lamellae under the fourth toe. Colour: Dorsally light brown to almost black with irregular lighter and darker flecks that usually coalesce to form a pair of broken middorsal stripes from the neck to the base of the tail. The parietals are often lighter in colour than the head and dorsum. The upper and lower labials are barred with light and dark streaks. Iris bright orange. An indistinct broken, darker dorsolateral line bordered above by a narrow lighter line is present from the eye and along the body and part of the tail. The tail is sometimes lighter in colour than the body. Laterally the body is flecked with light and dark markings which sometimes take the form of irregular striping. Ventrally grey to cream, darker spots usually present under the tail. The whole body has a strong opalescent sheen. Natural History Notes: A uncommon species. It is mainly diurnal and often moves about in the forest in open areas with ample shade, foraging for the small insects and their larvae that it feeds upon. It can also be found sheltering under leaf litter and fallen timber but does not appear to be a burrower, darting into the surrounding leaf litter if its cover is removed.

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ELEGANT FOREST SKINK Sphenomorphus concinnatus (Boulenger, 1887) Figs. 73, 74 Distribution: Endemic to the Solomon Islands: Bougainville, Shortland Islands, Fauro, Choiseul, Rob Roy, Vella lavella, Ranongga, Gizo, New Georgia, Tetepare, Vangunu, Isabel, Guadalcanal, Ngela, Malaita. Type locality: Fauro Island, Solomon Islands. Description: Average SVL 65mm, tail about 100% of SVL. Head moderate, snout quite short. Body fairly stout. Limbs well developed, overlapping when adpressed. A supranasal scale is usually absent but may be present in some specimens. Prefrontals in contact. A single anterior and posterior loreal (a small scale is often present above the anterior loreal, this could be regarded as a double anterior loreal). Frontal kite-shaped, about 2x as long as wide, longer than its distance from the snout, in contact with the first and second of 4 (very rarely 5) supraoculars. Lower eyelid scaly. No enlarged paired nuchals though some individuals may have 1 or 2 enlarged nuchals. Precloacals slightly enlarged. Midbody scales in 36 to 47 rows. 18 to 27 lamellae under the fourth toe. Colour: Dorsally golden brown with a more or less regular series of darker flecks. Usually a darker brown laterally with whitish spots that may form vague vertical bars. Many specimens have a black line running from the eye and onto the neck. On some islands these lizards have a large black spot on the side of the neck behind the ear, while on other islands this distinctive marking is absent (cf. figs. 73 and 74). Ventrally yellowish to dull orangebrown, this colour often sharply differentiated from the grey of the throat area. The tail is coloured similarly to the body. Natural History Notes: This common species is mostly diurnal though at times crepuscular. Greer and Parker (1967b) report S. concinnatus as being normally crepuscular on Bougainville. Elsewhere in the Solomons, particularly on Guadalcanal and Ngela where I have observed S. concinnatus, it is largely diurnal in its habits. It is terrestrial and occurs in forests and semi-cleared areas provided there is ample canopy cover and is often encountered sunning itself in patches of weak sunlight. It forages in the leaf litter for the small insects and spiders that if feeds upon. It is quite an active lizard and rapidly darts into the leaf litter if any attempt is made to approach it. It gives birth to 1 to 3 young. SKINKS, Family SCINCIDAE

79

CRANE’S SKINK Sphenomorphus cranei Schmidt, 1932 Figs. 75, 76, 77 Distribution: Endemic to the Solomon Islands: Bougainville, Shortland Islands, Vella lavella, New Georgia, Tetepare, Vangunu, Isabel, Guadalcanal, Ngela, Malaita. Type locality: Isabel Island, Solomon Islands. Description: Average SVL 60mm, tail about 150% of SVL. Head moderate. Body elongated. Limbs moderate, just meeting when adpressed. Supranasals absent. Prefrontals in contact or narrowly separated. A single anterior and posterior loreal. Frontal about 2x as long as wide, in contact with the first and second of 4 supraoculars. Lower eyelid scaly. Enlarged paired nuchals usually present. Precloacals slightly enlarged. Midbody scales in 32 to 41 rows. 20 to 29 lamellae under the fourth toe. Colour: Dorsally light or dark brown to almost black with light and dark flecking. Laterally similar to the dorsal colouration, though there is often an irregular pattern of light and dark bars that is more pronounced on the neck and anterior body. The lips are usually barred with black and white. The anterior venter is yellowish, becoming orange-red posteriorly, this colour extending to the underside of the tail. In some juveniles the posterior body and tail are bright orange-red. Natural History Notes: An uncommon species of secretive habits. It is sometimes diurnally active in heavily shaded areas otherwise it tends to a crepuscular existence. It is fairly dependent upon moisture and cool conditions and is usually found in the vicinity of forest streams, sheltering in holes in the bank or under rocks at the sides of these streams. On Ngela it is often encountered in association with Tribolonotus blanchardi, living under stones in moist stream beds. Sphenomorphus fragosus Greer and Parker, 1967 Fig. 78 Distribution: Endemic to the Solomon Islands: Bougainville. Type locality: Lake Loloru, Bougainville Island. Description: Average SVL around 60mm. Head moderate, snout short. Body a little stout. Limbs well developed, overlapping when adpressed. Head scales fragmented. Supranasals present, prefrontals separated by 1 or

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more small scales which also separate the frontal from the frontonasal. Two anterior and posterior loreals. The frontal is in contact with the first, second and third of 5 supraoculars. Lower eyelid scaly. Enlarged paired nuchals absent. Midbody scales in 42 to 46 rows. 18 to 21 lamellae under the fourth toe. Colour: Dorsally brown with transversely arranged darker spots. A black dorsolateral line is present. Laterally grey-brown, sometimes with lighter flecks. Ventrally yellow with or without darker spotting. The tail is marked similarly to the body. Natural History Notes: A poorly known species. Essentially montane and probably cryptozoic. Sphenomorphus solomonis (Boulenger, 1887) Figs. 79, 80 This species is almost certainly composite, not only over its entire range, but probably also within the Solomons populations (Glenn Shea, pers. comm). Distribution: Maluku Islands (Moluccas) in Indonesia, through mainland New Guinea and its adjacent island archipelagos to the Solomons. In the Solomons: Buka, Bougainville, Shortland, Fauro, Choiseul, New Georgia, Isabel, Guadalcanal, Savo, Ngela, Malaita, Ugi, Makira, Santa Cruz, Taumako, Reef Islands. Type locality: Fauro Island, Solomon Islands. Description: Average SVL 50mm, tail about 120% of SVL. Head small, snout short. Body elongated. Limbs poorly developed, not meeting when adpressed. Digits very small. Supranasals absent. Prefrontals separated. A single anterior and posterior loreal. Frontal about 2x as long as wide, longer than its distance from the snout, in contact with the first and second of 4 supraoculars. Lower eyelid scaly. Enlarged nuchals present. Precloacals slightly enlarged. Midbody scales in 22 to 30 rows. 11 to 18 lamellae under the fourth toe. Colour: Dorsally and laterally dark brown with a peppering of fine or coarse black and whitish flecks. This colour and pattern usually extends to the tail. Yellowish ventrally, often with a scattering of darker spots. All the body scales are very smooth and glossy. Natural History Notes: A common cryptozoic species found in forests and sometimes also in plantations and domestic gardens. It lives under fallen timber and mounds of ground debris in moist conditions. On SKINKS, Family SCINCIDAE

81

Guadalcanal it is often found in company with Tribolonotus schmidti. It is crepuscular and nocturnal, foraging in leaf litter for the small arthropods that it feeds upon. It is oviparous, laying up to 3 eggs. TANNER’S SKINK Sphenomorphus tanneri Greer and Parker, 1967 Fig. 81 Distribution: Endemic to the Bismarck Archipelago (New Britain, New Ireland) and the Solomon Islands: Bougainville, Shortland, Choiseul. Type locality: Kunua, Bougainville Island. Description: Average SVL 35mm, tail about 110% of SVL. Head small, snout obtusely conical. Limbs small, not meeting when adpressed. Supranasals absent. Prefrontals in contact. A single anterior and posterior loreal. Frontal only slightly longer than wide, in contact with the first and second of 4 supraoculars. Lower eyelid scaly. No enlarged nuchals. Enlarged precloacals present. Midbody scales in 28 to 32 rows. 13 to 20 lamellae under the fourth toe. Colour: Dorsally dark brown to almost black with small pale spots on the body and tail. Lighter spots are present dorsolaterally, often forming a distinct line. Laterally similar to the dorsal colouration though lighter. The gular area is brown to black, fading to yellowish ventrally, venter without markings. The area around the vent and underside of the tail is dull red. Natural History Notes: Quite common on Bougainville, apparently uncommon elsewhere in its range. A cryptozoic species preferring moist conditions, usually under rotting vegetation and fallen timber. It is oviparous, laying 2 eggs. Sphenomorphus taylori Burt, 1930 Fig. 82 Distribution: Endemic to the Solomon Islands: Bougainville. Type locality: Bougainville Island. Description: Average SVL 150mm, tail about 130% of SVL. Body elongated. Head shields fragmented, subject to variation. Supranasals usually present. Prefrontals separated by one or more small scales. 2 or more anterior loreals. Frontal usually entire or may be divided, longer than

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its distance from the snout, in contact with the first, second and third of 5 to 7 supraoculars. Interparietal distinct. Enlarged nuchals usually absent. Midbody scales in 53 to 60 rows. 27 to 35 lamellae under the fourth toe. Colour: Dark brown dorsally with narrow yellowish transverse bands which are more distinct laterally. Ventrally yellowish brown, often with darker spotting. Natural History Notes: A rare and poorly known species. Essentially montane and probably cryptozoic. Sphenomorphus transversus Greer and Parker, 1971 Fig. 83 Distribution: Endemic to the Solomon Islands, known from a single specimen from Bougainville (fig. 83). Type locality: Vicinity of Kunua, Bougainville Island. Description: SVL 68mm, tail 92mm. Head moderate, snout pointed. Limbs well developed, overlapping when adpressed. Supranasals absent. Prefrontals barely separated. Frontal longer than wide, longer than its distance from snout, in contact with first, second and third of 5 supraoculars. Lower eyelid scaly. Interparietal distinct. Enlarged nuchals absent. Midbody scales in 36 rows. 28 to 29 lamellae under the fourth toe. Colour: Dorsally light olive with a series of darker transverse bands from the neck to the base of the tail. A dark stripe runs through the eye. Ventrally the body and tail are yellow. Natural History Notes: Greer and Parker (1971) report the unique specimen as collected “under a decaying log on the steep side of a montane river valley covered with tall primary forest.” Sphenomorphus woodfordi (Boulenger, 1887) Fig. 84 Distribution: Endemic to the Solomon Islands: Bougainville, Shortland Island, Fauro, Choiseul, Ngela, Makira, Ugi. Type locality: Fauro Island, Solomon Islands. Description: Average SVL 120mm, tail about 120% of SVL. Head small, snout short. Body somewhat elongated. Limbs short, just meeting when adpressed. Supranasals absent. Prefrontals usually in contact. A single SKINKS, Family SCINCIDAE

83

anterior and posterior loreal. Frontal about 1.5x as long as wide, longer than its distance from the snout, in contact with the first and second of 4 (rarely 5) supraoculars. Lower eyelid scaly. Interparietal a little reduced in size. No enlarged nuchal scales. Midbody scales in 33 to 38 rows. 15 to 20 lamellae under the fourth toe. Colour: Dorsally light to dark brown with irregular darker and lighter spots. The darker markings tend to be more distinct on the posterior body and laterally where they may form vague transverse bars. Ventrally creamy white to light yellow. The tail often has a fine peppering of grey spots laterally. The whole body has a strong metallic gloss. Natural History Notes: Apparently reasonably common on Bougainville, uncommon elsewhere in its range. It lives in forests where it is largely diurnal though at times crepuscular. It forages for its insect food in the leaf litter on the forest floor and shelters under fallen timber and other ground debris. On Ngela I have seen S. woodfordi take refuge down the burrows of land crabs; it is possible that it utilizes these presumably empty burrows as a normal nocturnal retreat. Sphenomorphus sp. Fig. 85 In the first edition of this book (McCoy, 1980) I referred this species to Sphenomorphus undulatus which is widely distributed through New Guinea and is probably a composite species. However Glenn Shea (pers. comm.) believes the Solomons form, though closely related to Sphenomorphus undulatus, to be distinct from it and for the present should be treated as an undescribed species. Distribution: Apparently restricted the eastern Solomon Islands of Makira, the Olu Malau Group and Ugi. Description: Average SVL 40mm, tail about 150% of SVL. Head moderate. Body elongate. Limbs short, just meeting when adpressed. Supranasals absent. Prefrontals broadly in contact. A single anterior and posterior loreal. Frontal longer than wide, longer than its distance from the snout, in contact with the first and second of 4 supraoculars. Interparietal slightly reduced in size. Enlarged nuchals present. Midbody scales in 26 to 30 rows. 18 to 23 lamellae under the fourth toe. Colour: Dorsally light or dark brown with a series of lighter blotches which may be arranged into irregular transverse bands in some individuals. A

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broken, though distinct black line runs from the snout, through the eye and along the body in a dorsolateral position and continues onto the tail. Laterally with a scattering of large whitish spots and finer darker flecks. Ventrally yellow-orange to red in the region of the vent and underside of the tail. Some specimens have a scattering of fine black spots ventrally. Natural History Notes: This cryptozoic species inhabits the loose earth and debris in and under fallen timber and is also a common inhabitant of piles of husks in coconut plantations. It is mainly crepuscular though it is sometimes active diurnally in areas of heavy shade or when conditions are overcast. It shows a fair tolerance to dry conditions, often being found under rocks and timber where there is little moisture. On Makira it tends to inhabit the drier ridge tops rather than the wetter valleys. It is reasonably common in the eastern Solomons. Genus Tribolonotus Duméril and Bibron, 1839 Small to moderate sized lizards. Lower eyelid scaly. No supranasals or prefrontals. The frontoparietals may be distinct or fused. The head shields are quite rugose with longitudinal ridges. The body scales are carinate or spinose. Limbs reasonably well developed, pentadactyl. KEY TO THE SPECIES OF SOLOMON ISLANDS Tribolonotus 1. Enlarged vertebral scales in a double row: 2 – Enlarged vertebral scales in a single row: blanchardi 2. Males with palmar pores, average SVL less than 75mm: 3 – Males without palmar pores, average SVL more than 100mm: ponceleti 3. Enlarged vertebral scale rows contacting the parietal: schmidti – Enlarged vertebral scale rows not contacting the parietal: pseudoponceleti BLANCHARD’S CROCODILE SKINK Tribolonotus blanchardi Burt, 1930 Figs. 86, 87 Distribution: Endemic to the Solomon Islands: Bougainville, Choiseul, Isabel, Ngela. Type locality: Choiseul Island, Solomon Islands. SKINKS, Family SCINCIDAE

85

Description: Average SVL 35mm, tail about 160% of SVL. Head moderate, wedge-shaped, snout short. Body slender. Adpressed limbs overlap. Prefrontals absent. A single large loreal. Frontonasal large, longer than wide. Frontal reduced in size, smaller than the frontonasal. 4 supraoculars. Lower eyelid scaly. A single frontoparietal. Interparietal and parietals fused into a single shield. Mental small, followed by a large postmental and 2 pairs of chinshields. Head scales rugose, multicarinate. A cluster of enlarged spinose scales is present on each side of the head behind the parietal. Dorsal body scales small and strongly keeled. A single row of enlarged keeled vertebral scales runs from the neck to the base of the tail. An undulating row of enlarged scales is present along the sides of the body in a dorsolateral position. Ventrally with large keeled scales. A pair of enlarged keeled precloacals is present. Males with palmar and plantar pores and abdominal glands. 21 to 25 lamellae under the fourth toe. Colour: Colour and pattern highly variable. Dorsally light yellow to light or dark brown, this colour usually confined to within the vertebral area bordered by the undulating dorsolateral scale rows. Other specimens have a series of yellowish or greenish paired blotches along each side of the vertebral scale row. A light chevron shaped mark is normally present at the base of the tail. The lips are usually barred with black and whitish bands and the snout is often lighter in colour than the rest of the head. A short, broad white bar is usually present on the side of the head in front of the eye. The lateral and ventral colouration is mostly a uniform dark brown. The tail may be a uniform light or dark brown or it may be marked with broad light and dark bands. Natural History Notes: This species is quite common on Ngela at least, where it lives in very moist conditions in river valleys or under stones and piles of dead leaves and other debris in wet stream beds. On Bougainville it is essentially a montane lizard. I have seen T. blanchardi diurnally active on Ngela where it forages in and over debris at the sides of streams in heavily shaded areas in search of its food of small insects and their larvae. Although it generally moves fairly slowly, it is capable of bursts of rapid movement. If it is disturbed when sheltering under ground debris it either remains motionless or darts into the surrounding litter. It is oviparous, laying a single egg at a time.

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PONCELET’S CROCODILE SKINK Tribolonotus ponceleti Kinghorn, 1937 Figs. 88, 89, 90 Distribution: Endemic to the Solomon Islands: Bougainville, Shortland, Isabel. Type locality: Bougainville Island. There is some doubt as to whether this species actually occurs on Bougainville. Greer and Parker (1968a) point out that the species is absent in the extensive collections made on the island by Parker in the 1960s. Description: Average SVL 120mm, tail about 130% of SVL. Head quite large (proportionally smaller in the single known female specimen), snout short and angular. Adpressed limbs overlap. Prefrontals absent. A single large loreal is present. Frontonasal quite large. Frontal a little longer than wide, shorter than its distance from the snout. 4 supraoculars. 3 supralabials, the second very large. Lower eyelid scaly. Parietals and interparietal fused into a single shield. Mental small, followed by a larger postmental and 2 pairs of chinshields. All the head shields are rugose and multicarinate. Dorsal and caudal scales are all strongly spinose. A double row of enlarged keeled scales run from the neck to the tail in a vertebral position. The ventral scales are enlarged and strongly keeled. Males with 3 or 4 plantar pores and paired abdominal glands, no palmar pores. The only known adult female has a pair of abdominal glands, 1 plantar pore on the left foot and none on the right, no palmar pores. 25 to 28 lamellae under the fourth toe. Colour: Dorsally dark brown, the head and sides a lighter shade of brown. There are a series of lighter coloured irregular transverse bands from the neck, along the body and continuing onto the tail. Ventrally cream. A hatchling was dark brown dorsally, black on its lateral surfaces and tail, with a light creamy-yellow head, the transverse banding was white and more distinct than in the adults. As this lizard grew the banding faded and the head became darker. Natural History Notes: This species is very rare, currently known from fewer than 10 specimens, including an adult pair collected in April 1984 by local people near Harapa Village on Shortland Island. These latter two lizards were found together under a large log in undisturbed forest. At the time of collection the adult male was 122mm SVL, and the adult female was 112mm SVL. I maintained these lizards in captivity, and in March 1985 they produced SKINKS, Family SCINCIDAE

87

a male offspring which hatched from an egg (fig. 88). At hatching this lizard had a total length of 110mm (SVL 47mm, tail 63mm). In December 1986, when this lizard had almost reached adult size (SVL 98mm, tail 141mm), it was killed by the larger adult male. The adult male died several months later after developing an abscess on its jaw. The remaining female lived until 1994, presumably having reached the end of her normal lifespan. These lizards are now deposited in the Australian Museum and Bishop Museum. Tribolonotus ponceleti is a burrowing species and feeds on insects and spiders. It is completely nocturnal and by day shelters under fallen timber where it excavates tunnels to several entrances from a central chamber. The captive adult lizards - particularly the male - were noticeably aggressive, assuming a raised posture with open mouth and vigorously attempting to bite if disturbed in any way. This skink also vocalises; a sharp, high-pitched squeak that is used in conjunction with the open-mouthed threat display. WESTERN CROCODILE SKINK Tribolonotus pseudoponceleti Greer and Parker, 1968 Fig. 91 Distribution: Endemic to the Solomon Islands: Buka, Bougainville, Choiseul. Type locality: Kunua, Bougainville Island, Papua New Guinea. Description: Average SVL 70mm, tail about 120% of SVL. Head fairly large, wedge-shaped. Adpressed limbs overlap. Nostril in a single nasal. Prefrontals absent. Frontal smaller than the frontonasal. 4 supraoculars. Lower eyelid scaly. Parietals and interparietal fused into a single shield. Head shields rugose and multicarinate. Body scales small and granular or spinose. A double row of enlarged keeled scales runs from a little anterior to the forelimbs to the base of the tail in a vertebral position. Ventrally with enlarged keeled scales. The tail is covered with keeled scales forming annuli. Two enlarged, keeled precloacal scales are present. Males with palmar pores, both sexes with abdominal glands. 20 to 29 lamellae under the fourth toe. Colour: Dorsally a shade of dark brown to black. The head is usually black. Laterally and ventrally a medium brown. A series of large pale tan blotches are often present laterally. Specimens from Buka are generally lighter in colour than those from Bougainville (Greer and Parker 1968b). Natural History Notes: This lizard is very common on Bougainville and Choiseul where on the former island it occurs from coastal regions up to

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1200m elevation (Greer and Parker 1968b). Although it is sometimes found in semi-cleared areas, such as under the piles of discarded husks in coconut plantations, it is generally a forest dweller where it shelters under fallen timber and other debris on the forest floor. This species is oviparous, laying a single egg at a time. SCHMIDT’S CROCODILE SKINK Tribolonotus schmidti Burt, 1930 Figs. 92, 93 Distribution: Endemic to the Solomon Islands: Guadalcanal. Type locality: Beagle Island (now known as Marapa Island) in Marau Sound (east Guadalcanal), Solomon Islands. Burt and Burt (1932) record this species from Bougainville although it almost certainly does not occur there, Burt and Burt’s specimen is probably referable to T. pseudoponceleti. Description: Average SVL 35mm, tail about 100% of SVL. Head large (proportionally smaller in females), wedge-shaped, snout short. Limbs moderate, meeting when adpressed. Nostril in a single large nasal. Prefrontals absent. A single large loreal. Frontal smaller than the frontonasal, shorter than its distance from the snout. 4 supraoculars. Lower eyelid scaly. Frontoparietal wider than the frontal though shorter. Parietals and interparietal fused into a single shield. Mental large, followed by a pair of equally large postmentals. Head shields rugose, multicarinate. Dorsally the body is covered with small keeled or spinose scales interspersed with larger spinose scales that usually tend to form irregular longitudinal rows. A double row of enlarged keeled scales runs from the parietal to the tail. Tail covered with large, heavily keeled scales. A pair of enlarged keeled precloacals is present. Males with palmar and plantar pores and both sexes with abdominal glands. 15 to 25 lamellae under the fourth toe. Colour: Usually a uniform dark brown dorsally and laterally and ventrally light brown to dark reddish brown. On many specimens the middorsal area is lighter in colour than the rest of the dorsum. A light chevron mark is sometimes present at the base of the tail, and in most specimens the lips are barred with a few narrow white streaks. Iris bright to dull orange. Natural History Notes: A very common species. It is cryptozoic in habit and generally lives in moist conditions under fallen timber in forest areas. One large fallen tree may shelter several individuals, often an adult male SKINKS, Family SCINCIDAE

89

and female and several juveniles, possibly indicating that at least in this species of Tribolonotus, long-term pair bonds are formed. When cover is removed they remain motionless or dart for shelter in the surrounding leaf litter. This lizard is quite dependent upon cool, moist conditions and in direct sunlight, or if held in the hand for too long, will succumb to overheating. I have never seen an individual moving in the open of its own accord. It is probable that it only leaves its shelter if extended dry periods compel it to seek moister surroundings. Captive T. schmidti sleep at night so that like T. blanchardi it is probably mostly diurnally active. It feeds mainly on small insects and their larvae, captive specimens feed readily on termites. Unlike other members of its genus which are oviparous, T. schmidti gives birth to a single young.

MONITOR LIZARDS, Family VARANIDAE A family of moderate to very large lizards occurring largely in tropical areas though some are to be found in temperate climates. They are distributed from Africa through Asia to New Guinea and some of the islands of the tropical western Pacific and to Australia where the majority of species occur. In Australia they are called goannas, elsewhere they are usually termed monitor lizards. The head is fairly long and slender and the tail is long and laterally compressed in many species. The body is covered with small juxtaposed scales. The limbs are well developed and strongly clawed and the tongue is long and bifid, retractile into a sheath at its base. Monitor lizards are diurnal and occupy a variety of habitats from deserts to forests and some are aquatic to a large extent. They are voracious predators, feeding on a variety of vertebrates and invertebrates. All species are oviparous.

Genus Varanus Merrem, 1820 See family for description. KEY TO THE SPECIES OF SOLOMON ISLANDS Varanus – Nuchal and dorsal scales smoothly rounded, dorsally with a pattern of more or less evenly distributed small spots: indicus

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MONITOR LIZARDS, Family VARANIDAE

– Nuchal and dorsal scales spinose, dorsally with small spots interspersed with a series of much larger spots which form distinct transverse bands: spinulosus PACIFIC MONITOR Varanus indicus (Daudin, 1802) Figs. 94, 95 Böhme, Kai and Ziegler (2002) have referred the Rennell Island population of this lizard to a new species, Varanus juxtindicus. However, one of the main characters upon which they largely base their diagnosis - the base of the tail being round in cross-section in juxtindicus (supposedly laterally compressed in indicus) - does not adequately differentiate the Rennell animals from the many other Solomons populations of V. indicus that I have observed, in which the base of the tail is normally more or less round in cross-section, though it may be a little laterally compressed in some individuals. While juxtindicus may prove to be a valid taxon, for the present time, in the absence of more detailed comparative studies of other populations of Solomons varanids from different islands, I regard V. juxtindicus as a synonym of V. indicus. Distribution: Caroline, Marshall and Mariana Islands, the Indonesian archipelago, Timor, New Guinea, northern Australia, Torres Strait, Solomon Islands. In the Solomons: Buka, Bougainville, Shortland, Choiseul, Taro Island, Vella lavella, Gizo, Kolombangara, New Georgia, Tetepare, Rendova, Vangunu; Isabel, Russell Islands, Guadalcanal, Savo, Ngela, Rennell, Malaita, Makira, Ugi, Olu Malau Group, Santa Ana. Type locality: Ambon, Maluku (Moluccas), Indonesia. Description: Average SVL 500mm, tail about 170% of SVL. Head large, snout elongate. Canthus rostralis distinct. Tongue long and bifid. The head is covered with irregular polygonal scales which are more uniform on the supraocular region where they are transversely enlarged. The nostril is situated on the side of the snout, about two-thirds the distance from the eye to the tip of the snout. The eye is large and situated midway between the nostril and the ear. Body scales juxtaposed and somewhat conical dorsally. Ventrally the scales are smooth and rectangular. The caudal scales are carinate and rectangular. The tail is normally more or less round in cross section at its base though for most of its length it is strongly compressed laterally with a caudal keel. The limbs are well developed and strongly clawed. MONITOR LIZARDS, Family VARANIDAE

91

Colour: Black dorsally with small yellow spots arranged evenly over the upper surfaces of the body. On the tail these spots are often arranged into ill-defined transverse bars. The tip of the snout and the lips are often an orange-pink, particularly in younger animals. Ventrally cream to yellow. Natural History Notes: This lizard is uncommon on the major islands, although on many of the smaller islands such as the Olu Malau group, very large populations occur. Prior to the Second World War it was apparently quite common on the northern coast of Guadalcanal, an area where it no longer occurs. It is likely that the introduction of the toxic toad Bufo marinus in the late 1930s has been responsible for the demise of the monitor on northern Guadalcanal and the other main islands where the toad also occurs as monitor lizards eating this toad invariably die in a short time. Solomon Islanders in rural areas eat this lizard, its fat bodies in particular being regarded as delicacies. Varanus indicus is an active diurnal lizard and is usually found in the vicinity of the sea or mangrove swamps where it forages for its food of crabs (grapsid crabs in the littoral zone are a major part of its diet), smaller lizards and snakes, rats, and occasionally birds and their eggs. It will also dig up the nests of turtles to feed on the eggs. Although it is largely terrestrial in its feeding habits, it is a very efficient climber and invariably takes shelter up the nearest tree when alarmed on the ground. Monitors living around the shore of Lake TeNggano on Rennell are largely aquatic. This monitor prefers semi-open areas to thick forests and is a common inhabitant of coconut plantations on the smaller islands. The eggs are laid under decaying vegetation on the ground or in the rotting timber of dead trees on or above the ground. The hatchling lizards feed upon insects and small skinks and geckos. ISABEL MONITOR Varanus spinulosus Sprackland, 1994 Fig. 96 This varanid was originally described by Mertens (1941) as a Solomons endemic subspecies, Varanus indicus spinulosus. Distribution: Endemic to the Solomon Islands: San Jorge and Isabel. Type locality: San Jorge, Solomon Islands. Description: Average SVL 320mm, tail about 175% of SVL. Nuchal and dorsal scales spinose. Snout relatively short (when compared to V.

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MONITOR LIZARDS, Family VARANIDAE

indicus), and sharply tapered. Ventral scales rectangular and carinate. Tongue long and bifid, uniformly pink in colour. The tail is compressed laterally, forming a distinct caudal keel distally. Limbs are well developed and strongly clawed. Colour: Black to dark brown dorsally with a regular scattering of small yellowish spots. From the neck to the level of the hindlimbs a series of large yellow or yellow-green spots form regular transverse bands. The head is uniformly dark brown to almost black, often with a sparse scattering of small yellow spots. Tail with narrow, distinct bands for most of its length. Ventrally dark grey to brown with sparse lighter mottling. Natural History Notes: Relatively common on both San Jorge and the south-eastern end of Isabel. The largest populations occur in the extensive mangrove areas on northeast San Jorge and on the adjacent coast of Isabel. The ecology of V. spinulosus is essentially similar to that of V. indicus.

SNAKES, Order SQUAMATA, Suborder SERPENTES KEY TO THE FAMILIES OF SOLOMON ISLANDS SNAKES 1. Tail more or less rounded in cross section: 3 – Tail laterally compressed, paddle-shaped: 2 2. Ventral scales more than 3x as wide as the surrounding body scales: Elapidae (sea kraits) – Ventral scales only slightly wider than surrounding body scales: Elapidae (sea snakes) 3. Body scales smooth, or if keeled, then ventral scales distinct: 4 – Body scales keeled or spinose, distinct ventral scales absent: Acrochordidae 4. Body scales smooth: 5 – Body scales with distinct keels: Boidae 5. Eyes well developed, ventral scales distinct: 6 – Eyes vestigial, only discernible as small dark spots on the head, ventral scales absent: Typhlopidae MONITOR LIZARDS, Family VARANIDAE

93

6. Loreal scales present, diameter of the eye greater than its distance from the mouth: Colubridae – Loreal scales absent, diameter of the eye the same as or less than, its distance from the mouth: Elapidae (terrestrial elapids)

BLIND SNAKES, Family TYPHLOPIDAE A family of small, nocturnal burrowing snakes occurring widely in tropical and temperate regions around the world. All are alike in appearance and general ecology. The body scales are very smooth and glossy and the tail is short. Most species have a terminal spine on their tail that is used as an aid to progression through the ground. The head is not distinct from the neck and the eyes are vestigial. The mouth is situated well under the head. They have no venom apparatus. They are all oviparous and feed on termites, ants and their eggs and larvae. In the Solomons blind snakes mostly live in forested areas, though sometimes can be found in domestic gardens, usually in moist conditions in and under rotting fallen timber and in loose soil. They are occasionally found moving over the ground at night. Because of their secretive habits, these small snakes are rarely encountered and their relative abundance in the Solomons is not known. KEY TO THE GENERA OF SOLOMON ISLANDS BLIND SNAKES – Midbody scales in 26 or more rows: Acutotyphlops – Midbody scales in 24 or fewer rows: Ramphotyphlops

Rostral

Eye Preocular

Nasal

Head scales of a typhlopid snake (Ramphotyphlops becki)

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BLIND SNAKES, Family TYPHLOPIDAE

Genus Acutotyphlops Wallach, 1995 Lower jaw V-shaped. Two or more subocular scales. The frontorostral bordered by a pair of prefrontals. 5 or more infralabials. Midbody scales in 26 to 36 rows. KEY TO THE SPECIES OF SOLOMON ISLANDS Acutotyphlops 1. Snout rounded when viewed from above: solomonis – Snout pointed when viewed from above: 2 2. Midbody scales in 26-28 rows: infralabialis – Midbody scales in 30-35 rows: kunuaensis RED BLIND SNAKE Acutotyphlops infralabialis (Waite, 1918) Fig. 97 Distribution:. Bougainville, New Georgia, Guadalcanal, Ngela, Malaita. Type locality: Malaita, Solomon Islands. Description: Average total length 250mm, grows to around 350mm. The snout is wedge-shaped when viewed dorsally and laterally. 26 to 28 scale rows at midbody. Colour: Pinkish brown to red above. Yellowish ventrally. The edges of the scales are pale, giving an overall net-like pattern. Natural History Notes: See familial description for general ecological remarks. KUNUA BLIND SNAKE Acutotyphlops kunuaensis Wallach, 1995 Distribution: Endemic to Bougainville. Type locality: Kunua, Bougainville, Papua New Guinea. Description: Total length up to 300mm. Snout pointed when viewed dorsally and laterally. Midbody scales in 30 to 36 rows. Colour: Dark brown dorsally, sharply differentiated from the lighter yellowish ventral area. Natural History Notes: See familial description for general ecological remarks. BLIND SNAKES, Family TYPHLOPIDAE

95

Acutotyphlops solomonis (Parker, 1939) Distribution: Known only from the Milne Bay area on mainland Papua New Guinea and several localities on Bougainville. Type locality: Bougainville, Papua New Guinea. Description: Total length around 350mm, grows to over 450mm. Snout rounded when viewed dorsally, pointed when viewed laterally. Midbody scales in 29 to 34 rows. Colour: Uniformly dark brown dorsally and lighter yellowish-tan ventrally. Natural History Notes: See familial description for general ecological remarks.

Genus Ramphotyphlops Fitzinger, 1843 This genus is distinguished from other genera in its family by differences in hemipenial morphology. KEY TO THE SPECIES OF SOLOMON ISLANDS Ramphotyphlops 1. Midbody scales in 20 or more rows: 2 – Midbody scales in 18 rows: mansuetus 2. Nasal cleft contacting the preocular: braminus – Nasal cleft not in contact with the preocular: 3 3. Nasal cleft contacting the first supralabial: angusticeps – Nasal cleft contacting the second supralabial: 4 4. Snout rounded when viewed dorsally and laterally: depressus – Snout rounded when viewed dorsally, wedge-shaped when viewed laterally: becki ARBOREAL BLIND SNAKE Ramphotyphlops angusticeps (Peters, 1878) Fig. 98 Distribution: New Caledonia, Solomon Islands. In the Solomons it is known from Guadalcanal, Malaita, Makira, Malaupaina (Olu Malau Group), Rennell. Type locality: New Caledonia.

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BLIND SNAKES, Family TYPHLOPIDAE

Description: Average total length 350mm. Body very long and slender. Snout prominent, slightly hooked. Rostral with a distinct transverse keel. The nasal cleft contacts the first supralabial. Midbody scales in 20 rows. Colour: Light or dark brown dorsally, slightly paler ventrally. The mouth and vent areas are often whitish. Natural History Notes: This species appears to be partially if not completely arboreal. Of the dozen or so specimens I have found, all have been either sheltering diurnally in the collected detritus in hollows in forest trees or actively foraging nocturnally through foliage up to 4 metres above the ground. BECK’S BLIND SNAKE Ramphotyphlops becki (Tanner, 1948) Fig. 99 This snake was originally described from a single specimen from Guadalcanal as Typhlops becki by Tanner (1948). This specimen was later referred to Ramphotyphlops (then Typhlina) willeyi (a Loyalty Islands species) by McDowell (1974). Shea and Wallach (2000), with access to additional museum specimens, resurrected the species from the synonymy of willeyi. Distribution: Endemic to Solomon Islands: Guadalcanal. Type locality: Guadalcanal, Solomon Islands. Description: Average SVL about 140mm (males up to 128mm SVL, females to 148.5mm [Shea and Wallach, 2000]). Snout depressed, wedgeshaped when viewed laterally, rounded when viewed dorsally. Rostral almost reaching the level of the eyes, about a quarter the width of the head. Nasal almost divided by a cleft which contacts the second supralabial. Midbody scales in 20 rows. Colour: Dark brown above, lighter brown on the head. Ventrally light brown. There are often pale yellowish areas around the snout and underside of the tail. Natural History Notes: This snake has been found on Guadalcanal up to 2400m elevation (Shea and Wallach, 2000). See familial description for general ecological remarks.

BLIND SNAKES, Family TYPHLOPIDAE

97

FLOWERPOT SNAKE Ramphotyphlops braminus (Daudin, 1803) Fig. 100 Distribution: This wide-ranging parthenogenetic species has been recorded from southern Africa, Saudi Arabia, Iran, Madagascar, Reunion Island, Pakistan, India, southern China, Hong Kong, Burma, Thailand, Laos, Vietnam, Taiwan, Philippines, Okinawa, Malaysia, Singapore, Guam, Belau, Saipan, Ponape, Indonesia, Christmas and Cocos (Keeling) Islands, New Guinea, the vicinity of Darwin and Torres Strait Islands in northern Australia, Solomon Islands, Hawai’i, western Mexico. Type locality: Bengal, India (Burt and Burt, 1932). In the Solomons it has to date been found on Bougainville, Guadalcanal and Tulagi. This species was not recorded from Papua New Guinea prior to the Second World War (McDowell, 1974), likely indicating recent human introduction. It is similarly a recent introduction to the Solomons. It most likely occurs on other islands of the Solomons, but is rarely noticed because of its small size and secretive habits. Description: Average total length 130mm. Snout rounded when viewed dorsally and laterally. Rostral without a keel, about a quarter to a third the width of the head, wider dorsally than at the snout, not reaching the level of the eyes. Nasal cleft dividing the nasal, contacting the preocular. Midbody scales in 20 rows. Colour: Dark purplish brown to almost black above, lighter ventrally. There are rows of whitish glands in the sutures between the head scales. Natural History Notes: This widespread species generally lives in close association with man over much of its range and has probably been introduced by man to many areas where it occurs. It was possibly introduced into the Solomons during World War II when it could well have been secreted as eggs or mature specimens amongst the large amounts of military equipment brought into these islands at that time. All known specimens are female, the species reproducing by parthenogenesis. See familial description for general ecological remarks.

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BLIND SNAKES, Family TYPHLOPIDAE

Ramphotyphlops depressus (Peters, 1865) Figs. 101, 102 Distribution: Papua New Guinea (Manus Island and the islands of the Bismarck Archipelago), Solomon Islands. In the Solomons it is known from Bougainville, Shortland, Vella lavella, Ranongga, Kolombangara, New Georgia, Isabel, Guadalcanal, Ngela, Malaita, Olu Malau group and Makira. Type locality: Duke of York Island, Bismarck Archipelago, Papua New Guinea. Description: Average total length about 200mm. Snout rounded. Rostral without a keel, about a third the width of the head, nearly reaching the level of the eyes. The nostril is in a large nasal which is usually completely divided by the nasal cleft. The nasal cleft contacts the second supralabial. Midbody scales in 22 to 24 rows. Colour: Light to dark yellowish-brown to dull orange above. Ventrally a lighter brown or tan. Natural History Notes: This is the most commonly encountered species of blind snake in forest areas in the Solomons. See familial description for general ecological remarks. Ramphotyphlops mansuetus Barbour (1921) Distribution: This Solomons endemic species is currently known from two specimens; one from Makira, the other from Bougainville. Given this disparate distribution it is likely the species also occurs on the intervening islands. The holotype was collected at “Keri Keri” (Kira Kira, now the provincial capital) on Makira and described as Typhlops cumingii mansuetus by Barbour (1921) and later referred to Typhlina affinis by McDowell (1974) before being assigned separate specific status by Wallach (2003). Type locality: Makira, Solomon Islands. Description: The two known specimens are females, the larger is 159mm total length (Wallach, 2003) and the smaller holotype is 150mm (McDowell, 1974). Snout short and bluntly rounded from above. Rostral very wide, almost half the width of the head. The nasal is the largest head scale and is completely divided by a cleft which contacts the second supralabial. The eyes are particularly small and only discernable as black dots at the junction of the ocular, preocular and supraocular scales. Midbody scales in 18 rows. BLIND SNAKES, Family TYPHLOPIDAE

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Colour: The following is McDowell’s (1974) description of the colour of the preserved holotype: “Color brown, with large head scutes and mouth region yellowish; anteriorly, scales with pale (glandular?) bases, but posteriorly scales with black margins.” Natural History Notes: See familial description for general ecological remarks.

BOAS, Family BOIDAE Boas are mainly New World snakes (though the genus Candoia occurs from the Indonesian archipelago to the islands of the western South Pacific). Boas tend to be largely tropical in their distribution, though some species occur in temperate regions. Although some boas are fairly small, the family includes some of the largest snakes in the world such as the South American Anaconda. They feed on mammals, birds and other reptiles. Most species are nocturnal though some will move about during the day. According to species they are either arboreal, terrestrial or partly aquatic. Boas are live-bearers.

Genus Candoia Gray, 1842 Head distinct from the neck. Snout obliquely truncate. Canthus rostralis prominent. Body slightly compressed. All the dorsal body scales are strongly carinate. Subcaudals entire. KEY TO THE SPECIES OF SOLOMON ISLANDS Candoia 1. Supralabials separated from the eye by subocular scales: 2 – Supralabials entering the eye: paulsoni 2. Body stout, keels on body scales forming oblique rows: aspera – Body fairly slender, keels on body scales parallel to the long axis of the body: bibroni

100

BOAS, Family BOIDAE

NEW GUINEA GROUND BOA Candoia aspera (Günther, 1877) Fig. 103 Distribution: New Guinea and adjacent island archipelagos. Type locality: Duke of York Island, Bismarck Archipelago, Papua New Guinea. Its occurrence in the Solomons is based on a single specimen in the Australian Museum (R6994) reported by Kinghorn (1928) to have come from Bougainville. I have examined this specimen, and while its identity is certain, there is some doubt as to its provenance as the species is absent in the extensive Bougainville collections made by Fred Parker in the 1960s. However, in the specimen register at the Australian Museum, this snake is listed as having been received as part of a collection of reptiles, all of which are typical Bougainville species. I therefore believe its occurrence on Bougainville remains a possibility. Description: Average total length about 750mm. Head large, distinct from the neck. Body short, stout. Rostral wider than deep, not visible from above. Eye separated from supralabials by a row of suboculars. Head covered with small, irregular carinate scales. Body scales strongly carinate, the keels forming oblique rows. Midbody scales in 30 to 45 rows. Ventrals 127 to 153. Anal entire. Subcaudals 11 to 22, entire. Colour: Dorsal ground colour reddish brown with irregular light yellowbrown patterning edged with black. Ventrally yellowish, with or without dark spots. Natural History Notes: A terrestrial species, usually found under or amongst leaf litter and other ground debris. It feeds on lizards and small ground-dwelling mammals. SOUTH PACIFIC TREE BOA Candoia bibroni (Hombron and Jacquinot, 1853) Figs. 104, 105, 106, 107 Distribution: Solomon Islands, Vanuatu, Loyalty Islands, Fiji Islands, Tokelau Islands, Wallis and Futuna, the Samoas, Tonga. In the Solomons: Rennell, Makira, Ugi, Olu Malau, Santa Ana, Santa Cruz, Reef Islands, Vanikoro, Utupua. Type locality: Viti Levu, Fiji. Description: Average total length 1.25m, sometimes reaches 2m. Head large, distinct from the neck, snout obliquely truncate. Body fairly slender. BOAS, Family BOIDAE

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Rostral wider than deep, not visible from above. The head is covered with small scales which possess a short, undeveloped keel. Diameter of eye greater than its distance from mouth. 10 to 15 supralabials separated from the eye by a row of suboculars. Body scales keeled, in 32 to 42 rows at midbody. 211 to 276 ventrals. Anal entire. 40 to 66 subcaudals, mostly entire though 1 or 2 may be divided. Colour: The colouration is quite variable. The commonest colour form is a grey ground colour above with irregular lighter and darker blotches, sometimes forming a variegated pattern. This patterning is usually more distinct on the posterior body and tail. Other colour forms are brick red with the same markings as above while other specimens may be dull or bright yellow with or without other markings. Ventrally cream to white with or without darker markings. Natural History Notes: This is an arboreal species and in some areas such as the Reef Islands where it is abundant, it can be found in mangroves. Elsewhere it is a dweller in forests and sometimes in cultivated areas. Mainly nocturnal, it occasionally moves about during the day. Food consists of lizards, particularly geckos, and mammals such as rats and flying foxes, and it very occasionally takes roosting and nesting birds. It is a slow moving snake, normally fairly docile, though aggressive when provoked, striking repeatedly if any attempt is made to come near it. It gives birth to up to 20 or more young. SOLOMONS GROUND BOA Candoia paulsoni (Stull, 1956) Figs. 108, 109, 110, 111, 112 Smith et al (2001) regard several of the Solomons populations of this snake as distinct subspecies though these designations are not considered here. Distribution: Parts of the Indonesian Archipelago, northern and eastern Papua New Guinea and the eastern Admiralty Islands, Solomon Islands. It occurs on virtually all the islands of the Solomons and Santa Cruz group with the exceptions of Ontong Java, Duff Islands, Tikopia and Anuta. Type locality: Ugi, Solomon Islands. Description: Average total length around 750mm, it grows to over 1m; the animal shown in fig. 110 is almost 1.2m in length. Head large, distinct from the neck, snout obliquely truncate. Canthus rostralis angular. Body

102

BOAS, Family BOIDAE

stout. Tail short and prehensile. Rostral wider than deep, not visible from above. The head is covered with small keeled or partially keeled scales. Eye moderately large with a vertically elliptic pupil. 10 to 14 supralabials of which 2 enter the eye. Body scales are strongly keeled, in 32 to 43 rows at midbody. 167 to 202 ventrals. Anal entire. 36 to 51 subcaudals, all entire. Colour: The basic colour may be light cream or brown, grey, brick red to dark brown above with darker zigzag patterning along the vertebral line and a variegated pattern laterally. Patches of irregular red and white blotches are present at the edge of the ventral area. Ventrally grey to white, usually with a peppering of grey, red and black spots. The subcaudal area is often patterned with a vague black and white check. Natural History Notes: This species is the most widespread of the snakes in the Solomons. It occurs abundantly in a variety of habitats from forests to cultivated areas and human habitations. It sometimes coils itself up in the open, relying on its cryptic colouration for protection. Mainly nocturnal, it occasionally moves about during the day. It is basically terrestrial though it is an efficient climber and sometimes can be found in tree hollows. On the ground it lives among any suitable cover from leaf litter to rock piles, grass tussocks and under fallen timber. When young, it feeds upon small lizards, particularly skinks, and larger snakes feed upon larger lizards and rats. I have found C. paulsoni deep within limestone caves where it was presumably feeding on small bats and/or juvenile frogs of the genus Discodeles that also occurred there. This snake can be aggressive in temperament, striking repeatedly if provoked. Up to 20 or more young are produced at a time.

FILE SNAKES, Family ACROCHORDIDAE This monogeneric family is represented in tropical regions from India and Sri Lanka, through most of southeast Asia and the Indo-Australian Archipelago to Papua New Guinea, northern Australia and the Solomons. These non-venomous snakes are totally aquatic and occur in marine, brackish and freshwater conditions. Their skin is loose and baggy and covered with very small spinose scales which gives rise to their common name of file snakes. They have no enlarged ventral or subcaudal scales. They feed on a variety of freshwater and marine fish and give birth to young.

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Genus Acrochordus Hornstedt, 1787 Head covered with irregular juxtaposed scales. Body scales small and spinose. Enlarged ventrals absent. Tail short and prehensile. Head scarcely distinct from the neck. Body stout and a little compressed. Eyes minute. Nostrils on the top of the snout, close together and valvular. LITTLE FILE SNAKE Acrochordus granulatus (Schneider, 1799) Fig. 113 Distribution: India, Sri Lanka, Pakistan, Bangladesh, China (Hainan), Cambodia, Indonesia, Malaysia, Myanmar (Burma), Philippines, Thailand, Vietnam, Singapore, New Guinea and its island archipelagos, northern Australia, Solomon Islands. In the Solomons it has been recorded from Bougainville, Shortland, Isabel, Ngela, Guadalcanal, Malaita. It probably occurs on other islands of the group. Type locality: unknown. Description: Average total length about 700mm. Head small, scarcely distinct from neck. Eyes minute. Nostrils on the top of the snout, valvular. Body covered with loose baggy skin with very small spinose scales giving a file-like appearance and texture. 85 to 108 scale rows at midbody. No distinct ventral or subcaudal scales. A distinct longitudinal fold is present midventrally. Colour: Dark grey to black with numerous pale, broad vertical bars laterally. In juveniles these bars are continuous on the dorsum. Natural History Notes: This common nocturnal species is totally aquatic and occurs in river estuaries, mangrove swamps and reef flats. It is particularly abundant in mangrove areas in Mboli Passage on Ngela and in the extensive mangrove areas bordering Lau Lagoon on Malaita. It feeds primarily on sedentary fish such as blennies, gobies and particularly mudskippers. When feeding, it wraps itself around the fish, its spinose skin preventing the fish from slithering free, and swallows it alive. McDowell (1979) reports snail shell fragments in the stomach of a specimen from Isabel Island. It is not an active species, often lying motionless on the bottom in shallow water for long periods. It is a live-bearer.

104

FILE SNAKES, Family ACROCHORDIDAE

COLUBRID SNAKES, Family COLUBRIDAE In the Australia-New Guinea-Pacific region colubrid snakes are poorly represented, though this is the largest family of snakes. They occur in tropical and temperate regions around the world. Generally they tend to be slender, agile snakes, either terrestrial or arboreal while some are aquatic to a large extent. They may be either diurnal or nocturnal. Typically they feed on amphibians, though other reptiles, birds and mammals are also eaten. The aquatic species feed mainly on fish. The family includes both solid-toothed non-venomous snakes and rear-fanged venomous snakes. KEY TO THE SPECIES OF SOLOMON ISLANDS COLUBRID SNAKES – Midbody scales in 13 rows: Dendrelaphis salomonis – Midbody scales in 17 to 23 (usually 21) rows: Boiga irregularis

Genus Boiga Fitzinger, 1826 Rear-fanged venomous land snakes. Head large, very distinct from neck. Eye large with a vertically elliptic pupil. Loreal scale present. BROWN TREE SNAKE Boiga irregularis (Merrem, 1802) Figs. 114, 115 Distribution: Maluku (Moluccas), Sulawesi, New Guinea and adjacent archipelagos, Australia, Solomon Islands. There is also an introduced population on Guam which has exterminated a number of species of native birds on that island. In the Solomons: Bougainville, Shortland, Mono, Choiseul, Ranongga, Vella lavella, New Georgia, Tetepare, Vangunu, Isabel, Guadalcanal, Ngela, Malaita. Type locality: unknown. Description: Average total length about 1.5m, grows to over 2m. Head very large, distinct from neck. Body slender, laterally compressed. Rostral wider than deep, visible from above. Usually 1 (sometimes 2) preocular which narrowly contacts the frontal or is narrowly separated from it. 2 postoculars. The eye is very large, its diameter greater than its distance from the mouth. Frontal slightly longer than wide, as long as its distance

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from the snout. 8 to 10 supralabials. Midbody scales in 17 to 23 (usually 21) rows. 217 to 270 ventrals. Anal entire. 99 to 125 subcaudals, all divided. Colour: Dorsally light brown to khaki with darker indistinct wavy transverse bands which are more distinct on the anterior body. Ventrally yellow or greenish yellow. The iris is usually orange-yellow. Natural History Notes: This common species is nocturnal and arboreal though it occasionally forages on the ground at night. It shelters in hollows in trees and in the crowns of various palms during the day and sometimes can be found in the ceilings of houses. It can be an aggressive species, striking repeatedly if provoked. Although it is rear-fanged and venomous, it is not regarded as dangerous to humans. It feeds mainly on lizards particularly geckos - and roosting or nesting birds and occasionally rats. It will also eat the eggs of wild birds and domestic chickens. It is oviparous.

Genus Dendrelaphis Boulenger, 1890 Solid-toothed non-venomous snakes. Head elongate, distinct from neck. Eye large with a round pupil. Body slender. Midbody scales in 13 to 15 rows. Loreal scale present. SOLOMONS TREE SNAKE Dendrelaphis salomonis (Günther, 1872) Fig. 116 Distribution: Misima Island (Papua New Guinea), Solomon Islands. In the Solomons: Buka, Bougainville, Shortland, Fauro, Choiseul, Rob Roy, Vella lavella, Gizo, Ranongga, Rendova, Kolombangara, New Georgia, Tetepare, Vangunu, Isabel, Russell Islands, Guadalcanal, Ngela, Malaita, Makira, Ugi, Olu Malau, Santa Ana, Santa Cruz, Utupua. Type locality: Solomon Islands. Description: Average total length 1m. Head elongate, distinct from the neck. Body very slender. Canthus rostralis rounded. Rostral wider than deep, visible from above. Nasal usually divided. Frontal bell-shaped, about 2x as long as wide, as long as its distance from snout. Eye large, its diameter greater than its distance from mouth. Temporals 1+2 or 2+2. 7 to 9 supralabials. Midbody scales in 13 rows. 171 to 211 ventrals. Anal entire. 124 to 166 subcaudals, all divided.

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COLUBRID SNAKES, Family COLUBRIDAE

Colour: Dorsally greenish grey to light or dark brown. The anterior third of the body is often reddish and in many individuals indistinct vertical darker bars are present on the anterior body. A black streak runs from the snout, through the eye and onto the neck. The labials and the gular region are invariably a pale yellow. Ventrally grey, grey-green, yellow-green, yellow or cream. Natural History Notes: A very active, swift moving diurnal snake. It is quite common and occurs in both forests and cultivated areas where it is both terrestrial and arboreal in its habits. This species is oviparous and preys mainly on lizards and occasionally frogs, the prey being eaten alive. This snake has very keen eyesight and will raise its head high to intently scan its surroundings when searching for prey. If it detects the movement of a small skink for example, it will make immediate pursuit, and with its superior speed is usually successful.

ELAPID SNAKES, Family ELAPIDAE (Subfamily Hydrophiinae) A large family of front-fanged venomous snakes. They occur from Africa, through Asia and Australia to New Guinea, Solomon Islands and Fiji in the southwest Pacific, and North and South America. Elapid snakes may be either diurnal or nocturnal. They feed on a variety of vertebrates (a few of the small fossorial species will eat invertebrate prey such as earthworms) and are either egg-layers or live-bearers. The Solomons elapid snakes are all members of the subfamily Hydrophiinae and are either land-dwelling (referred to here as terrestrial elapids) or live-bearing marine snakes (sea snakes) or egg-laying amphibious marine snakes (sea kraits). Typically the bodies of terrestrial elapids are more or less round in cross-section, the tail is round and tapering, and the ventral scales are distinct. They may be either egg-layers or live-bearers. The sea snakes are exclusively marine and are live-bearers. The nostrils are valvular, the ventral scales are barely distinguishable from the surrounding body scales, the body is laterally compressed and the tail is distinctly paddle-shaped. Sea snakes are inhabitants of the tropical and subtropical Indian and Pacific Oceans, though one species, Pelamis platurus, enters temperate waters. They feed on a variety of marine fish and fish eggs. The sea kraits are members of the ELAPID SNAKES, Family ELAPIDAE

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genus Laticauda. Although they feed exclusively in the marine environment, sea kraits are partly terrestrial, normally spending part of their daily activity cycle ashore, sheltering in rocky crevices (usually diurnally) and one species, Laticauda colubrina, can often found in tree hollows several metres above the ground (personal observation and Ineich and Laboute, 2002). All species are oviparous (although the reproductive mode of Laticauda crockeri is currently unknown), laying their eggs on land. The ventral scales are distinct and the tail is paddle-shaped (but not as markedly compressed as in the sea snakes). Most species of sea kraits are marked with a distinctive dark and light banded pattern. Sea snakes and sea kraits have a salt gland under their tongue; salt is accumulated in this gland and excreted when the snake protrudes its tongue. KEY TO THE SPECIES OF SOLOMON ISLANDS TERRESTRIAL ELAPID SNAKES 1. Body with distinct black and white banded pattern: Loveridgelaps elapoides – Body without distinct black and white banded pattern: 2 2. Small size (average total length 300mm), 32-35 subcaudals, occurring only on Bougainville: Parapistocalamus hedigeri – Moderate size (average total length 750mm), 38-59 subcaudals, not occurring on Bougainville: Salomonelaps par

Genus Loveridgelaps McDowell, 1969 A monotypic genus. Head slightly depressed, scarcely distinct from the neck. Eyes very small. Nasal single or divided. Midbody scales in 17 rows. Anal entire. SOLOMONS BLACK-BANDED KRAIT Loveridgelaps elapoides (Boulenger, 1890) Fig. 117 Distribution: Endemic to the Solomon Islands: Shortland, Choiseul, Rob Roy, Vella lavella, Gizo, Isabel, Guadalcanal, Ngela, Malaita. Type locality: Florida Islands (Ngela Islands), Solomon Islands.

108

ELAPID SNAKES, Family ELAPIDAE

Description: Average total length about 800mm, grows to over 1m. Head small, scarcely distinct from neck. Eyes very small. Body moderate. Rostral broad. Nostril in a single (occasionally divided) nasal which contacts the preocular. Frontal as wide as long, wider than the supraoculars. 7 supralabials, the third and fourth entering the eye. 1 or 2 postoculars. Temporals 1+2. Midbody scales in 17 rows. 193 to 218 ventrals. Anal entire. 31 to 38 subcaudals, all divided. Colour: Dorsally black with a regular series of bright yellow bands along the vertebral line. This yellow colour soon fades in preservative and previous accounts of this species do not mention this most distinctive colouration. Laterally with white banding, this colour often separated from the yellow bands by 1 or 2 rows of black scales. The head is usually white with irregular black markings on the rostral, labials and orbits and occasionally on the occiput. Melanotic forms of this snake are known from Malaita. Natural History Notes: This is a very rare and poorly known endemic species and is not usually found outside the forest areas where it lives, usually in the vicinity of streams. It is cryptozoic in its habits, sheltering under leaf litter and fallen timber and tends to be crepuscular or nocturnal. It is known to feed on blind snakes, and on Guadalcanal I found a Loveridgelaps of approximately 1m in length that had swallowed a Salomonelaps par of equal length to itself. It is an inoffensive snake and makes no attempt to bite, even when handled. However, as the toxicity of the venom of Loveridgelaps is unknown, its bite should be regarded as potentially dangerous. On Malaita Island this snake is known as baekwa i tolo (“shark of the forest”) and is greatly feared. The reproductive mode of Loveridgelaps is unknown, but it is probably oviparous.

Genus Parapistocalamus Roux, 1934 Two maxillary fangs. No posterior maxillary teeth. The head is not distinct from the neck. The diameter of the eye is less than its distance from the mouth. The nostril between 2 nasals. The body scales are in 15 rows. Anal entire or divided. Subcaudals all divided.

ELAPID SNAKES, Family ELAPIDAE

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HEDIGER’S SNAKE Parapistocalamus hedigeri Roux, 1934 Fig. 118 Distribution: Endemic to the Solomon Islands: Bougainville. Type locality: vicinity of Buin, Bougainville Island. Description: Average total length around 300mm. Head not distinct from the neck. Rostral broader than deep, visible from above. Nostril in a divided nasal. Frontal slightly longer than wide, longer than its distance from the snout. A single preocular may be present or absent. A single postocular. Temporals 1+1 or 1+2. Midbody scales in 15 rows. 159 to 169 ventrals. Anal entire or divided. 32 to 35 subcaudals, all divided.O’Shea (2005) suggests that the dentition of this snake indicates it is not closely related to other Solomon or Australasian elapids. Colour: (in preservative) Dorsally dark brown. Lighter laterally. The head is coloured similarly to the dorsum though the parietals are often light in colour. Ventrally yellow to light brown. Natural History Notes: A very rare species and very little is known of its habits. Williams and Parker (1964) report it as living “in and under rotten logs and leaf mould.”

Genus Salomonelaps McDowell, 1969 A monotypic genus. Head slightly depressed, distinct from the neck. Eye moderate. Nasal single or divided. Midbody scales in 15 to 17 rows. Anal divided. SOLOMONS RED KRAIT Salomonelaps par (Boulenger, 1884) Figs. 119, 120, 121, 122, 123 Distribution: Endemic to the Solomon Islands: Buka (small island immediately adjacent to northern Bougainville, but has not been recorded from Bougainville), Shortland, Fauro, Choiseul, Vella lavella, Ranongga, Simbo, Gizo, Kolombangara, New Georgia, Rendova, Tetepare, Vangunu, Isabel, Savo, Guadalcanal, Ngela, Guadalcanal, Malaita. Type locality: Fauro Island, Solomon Islands.

110

ELAPID SNAKES, Family ELAPIDAE

Description: Average total length about 750mm, grows to over 1m. Head moderate, distinct from neck. Body a little stout. Rostral wider than deep, visible from above. Nasal usually divided, though sometimes single, but at least always grooved. 1 preocular and 2 postoculars. Frontal as wide as long. 7 supralabials, third and fourth entering eye. Temporals 1+2. Midbody scales in 15 to 17 rows. 158 to 180 ventrals. Anal divided. 38 to 59 subcaudals, may be entire or divided or partly entire and partly divided. Colour: Populations from different islands vary considerably in colour and pattern (see figs 119 to 123). Usually a deep red or reddish brown to dark brown to almost black dorsally. Darker transverse bands may be distinct, indistinct or absent. In animals from New Georgia the scales have black borders resulting in an overall reticulated pattern. Ventrally creamy white. Natural History Notes: A reasonably common species throughout its range. It is essentially diurnal, though very occasionally nocturnal, and quite active. It occurs in forested areas, usually in the vicinity of streams where it forages for its prey of frogs and lizards. On Guadalcanal and Ngela at least, frogs of the genera Ceratobatrachus and Platymantis form a large part of its diet. It is also known to feed on blind snakes. When threatened this snake will flatten its neck and anterior body, hissing loudly. A chewing movement of the jaws, characteristic of many elapid snakes, may also occur. Although not particularly aggressive, it will bite if sufficiently provoked, and people from Ngela Island claim to know of dog and human fatalities resulting from its bite. I was bitten by a Salomonelaps on Choiseul Island and experienced no ill effects, most likely due to that individual snake’s failure to envenomate. The reproductive mode of Salomonelaps is unknown, but it is probably oviparous. KEY TO THE GENERA OF SOLOMON ISLANDS SEA SNAKES – Mental groove present, head normal, not elongate, body more or less distinctly banded: Hydrophis – Mental groove absent, head elongated, no dark transverse banding on the body: Pelamis

Genus Hydrophis Sonnini and Latreille, 1802 Head shields enlarged, regular. Mental groove present. The ventrals are much reduced in size. Four or more maxillary teeth following the fang. A key ELAPID SNAKES, Family ELAPIDAE

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is not given here for the species of Hydrophis. This is because there is some doubt as to the correct identification of some of the specimens from the Solomons (see species accounts), and also because members of this genus are difficult to identify to species level on the basis of external characters. Hydrophis belcheri (Gray, 1849) Distribution: Philippines, Indonesian Archipelago, New Guinea, Solomon Islands, Kiribati, Fiji. It is known in the Solomons from a single specimen collected off Malaita. This specimen was originally referred to Hydrophis fasciatus atriceps by Slevin (1934). Brown and Leviton of the California Academy of Sciences examined this specimen on my behalf and have informed me that they believe it is an example of Hydrophis belcheri. Description: The following is a description of the specimen (CAS 72035) forwarded to me by Brown. “Total length of specimen 640mm; posterior three-quarters of body and tail strongly compressed laterally; head small, not distinct from neck; rostral obtusely pointed dorsally, slightly broader than deep; frontal about 1.5x as long as broad; its length about two-thirds the length of the parietals and nearly as long as its distance from the rostral; eye small, its diameter about 1.25x its distance from the mouth and slightly less than its distance from the nostril, one preocular and 2 postoculars; a single, large anterior temporal; 6 upper labials, third and fourth entering the eye; 7 lower labials with 4 small scales at the oral margin above third, fourth and fifth; 29 scales on the neck; 36 on the body at midbody and at the point of greatest depth, pointed posteriorly and marked by a single keel; 335 ventrals each marked by two keels; precloacals enlarged. Colour in preservative: a pattern of alternating, complete, dark brown and dirty white, transverse bands; the bands of about equal breadth ventrally but the dark bands broader dorsally; 29 dark bands between the head and the tail.” Natural History Notes: Snakes of the genus Hydrophis appear to be uncommon or at least rarely seen in the Solomons. The few that I have seen have been in deeper water (between 20 and 30 metres) over a sandy, rather than a coral bottom and usually close to the mouths of large rivers.

112

ELAPID SNAKES, Family ELAPIDAE

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1 Crocodylus porosus Moitaka, Papua New Guinea

2 Chelonia mydas Malaupaina Island, Olu Malau Islands COLOR PLATES 1/2

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3 Eretmochelys imbricata Tandai Point, Guadalcanal Island

4 Lepidochelys olivacea Wainone Bay, Makira Island REPTILES OF THE SOLOMON ISLANDS

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5 Hypsilurus macrolepis (male) Malaupaina Island, Olu Malau Islands

6 Hypsilurus macrolepis (male) Kira Kira, Makira Island COLOR PLATES 3/6

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7 Hypsilurus macrolepis (female) Malaupaina Island, Olu Malau Islands

8 Cyrtodactylus biordinis Mt Austen, Guadalcanal Island REPTILES OF THE SOLOMON ISLANDS

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9 Cyrtodactylus louisiadensis Koliai, Shortland Island

10 Cyrtodactylus louisiadensis Mt Austen, Guadalcanal Island COLOR PLATES 7/10

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11 Cyrtodactylus louisiadensis (juvenile) Rokera, Malaita Island

12 Gehyra mutilata Marau, Guadalcanal Island REPTILES OF THE SOLOMON ISLANDS

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13 Gehyra oceanica Mboromole, Ngela Sule Island

14 Gekko vittatus Mboromole, Ngela Sule Island COLOR PLATES 11/14

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15 Gekko vittatus Malaupaina Island, Olu Malau Islands

16 Hemidactylus frenatus Mbua Valley, Guadalcanal Island REPTILES OF THE SOLOMON ISLANDS

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17 Hemiphyllodactylus typus Mbua Valley, Guadalcanal Island

18 Lepidodactylus flaviocularis Mt Austen, Guadalcanal Island COLOR PLATES 15/18

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19 Lepidodactylus guppyi Mboromole, Ngela Sule Island

20 Lepidodactylus guppyi Mbua Valley, Guadalcanal Island REPTILES OF THE SOLOMON ISLANDS

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21 Lepidodactylus lugubris Mbua Valley, Guadalcanal Island

22 Lepidodactylus woodfordi Korovou, Shortland Island COLOR PLATES 19/22

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23 Nactus multicarinatus Mt Austen, Guadalcanal Island

24 Carlia mysi Faisi Island, Shortland Islands REPTILES OF THE SOLOMON ISLANDS

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25 Corucia zebrata Korovou, Shortland Island

26 Corucia zebrata Mbua Valley, Guadalcanal Island COLOR PLATES 23/26

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27 Corucia zebrata (neonate) Dala, Malaita Island

28 Cryptoblepharus poecilopleurus Te-Uhungango, Rennell Island REPTILES OF THE SOLOMON ISLANDS

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29 Emoia atrocostata Kukum, Guadalcanal Island

30 Emoia caeruleocauda (male) Munda, New Georgia Island COLOR PLATES 27/30

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31 Emoia caeruleocauda (male) Ringgi, Kolombangara

32 Emoia caeruleocauda (female) Munda, New Georgia Island REPTILES OF THE SOLOMON ISLANDS

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33 Emoia caeruleocauda (juvenile female) Korovou, Shortland Island

34 Emoia caeruleocauda (male) Kira Kira, Makira Island COLOR PLATES 31/34

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35 Emoia caeruleocauda (female) Kira Kira, Makira Island

36 Emoia cyanogaster Lata, Santa Cruz Island REPTILES OF THE SOLOMON ISLANDS

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37 Emoia cyanogaster Noro, New Georgia Island

38 Emoia cyanura Mbua Valley, Guadalcanal Island COLOR PLATES 35/38

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39 Emoia cyanura Korovou, Shortland Island

40 Emoia flavigularis Mboromole, Ngela Sule Island REPTILES OF THE SOLOMON ISLANDS

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41 Emoia impar Mele, Efate Island, Vanuatu

42 Emoia isolata Bellona Island COLOR PLATES 39/42

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43 Emoia jakati Teopasino, Bougainville Island

44 Emoia maculata Kira Kira, Makira Island REPTILES OF THE SOLOMON ISLANDS

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45 Emoia maculata Malaupaina Island, Olu Malau Islands

46 Emoia nigra Mbua Valley, Guadalcanal Island COLOR PLATES 43/46

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47 Emoia nigra Mboromole, Ngela Sule Island

48 Emoia nigra Malaupaina Island, Olu Malau Islands REPTILES OF THE SOLOMON ISLANDS

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49 Emoia pseudocyanura Korovou, Shortland Island

50 Emoia pseudocyanura Falamai, Mono Island COLOR PLATES 47/50

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51 Emoia pseudocyanura Mbua Valley, Guadalcanal Island

52 Emoia pseudocyanura Mboromole, Ngela Sule Island REPTILES OF THE SOLOMON ISLANDS

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53 Emoia pseudocyanura Arabala, Malaita Island

54 Emoia rennellensis Te-Nggano, Rennell Island COLOR PLATES 51/54

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55 Emoia rufilabialis (male) Lata, Santa Cruz Island

56 Emoia rufilabialis (female) Lata, Santa Cruz Island REPTILES OF THE SOLOMON ISLANDS

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57 Emoia sanfordi Mele, Efate Island, Vanuatu

58 Emoia schmidti Munda, New Georgia Island COLOR PLATES 55/58

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59 Emoia schmidti Ringgi, Kolombangara Island

60 Emoia taumakoensis (male) Taumako Island, Duff Islands REPTILES OF THE SOLOMON ISLANDS

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61 Emoia taumakoensis (female) Taumako Island, Duff Islands

62 Eugongylus albofasciolatus Teopasino, Bougainville Island COLOR PLATES 59/62

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63 Eugongylus albofasciolatus Aruligo, Guadalcanal Island

64 Eugongylus rufescens Hutuna, Rennell Island REPTILES OF THE SOLOMON ISLANDS

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65 Geomyersia glabra Mboromole, Ngela Sule Island

66 Lamprolepis smaragdina Arabala, Malaita Island COLOR PLATES 63/66

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67 Lamprolepis smaragdina Nukiki, Choiseul Island

68 Lipinia noctua Mbua Valley, Guadalcanal Island REPTILES OF THE SOLOMON ISLANDS

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69 Prasinohaema virens Mboromole, Ngela Sule Island

70 Prasinohaema virens Malaupaina Island, Olu Malau Islands COLOR PLATES 67/70

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71 “Sphenomorphus” bignelli Mt Austen, Guadalcanal Island

72 “Sphenomorphus” bignelli Mboromole, Ngela Sule Island REPTILES OF THE SOLOMON ISLANDS

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73 Sphenomorphus concinnatus Mt Austen, Guadalcanal Island

74 Sphenomorphus concinnatus Anuta Island, Ngela Islands COLOR PLATES 71/74

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75 Sphenomorphus cranei Mt Gallego, Guadalcanal Island

76 Sphenomorphus cranei Mboromole, Ngela Sule Island REPTILES OF THE SOLOMON ISLANDS

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77 Sphenomorphus cranei (juvenile) Koliai, Shortland Island

78 Sphenomorphus fragosus (paratype) Lake Loloru, Bougainville Island COLOR PLATES 75/78

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79 Sphenomorphus solomonis Teopasino, Bougainville Island

80 Sphenomorphus solomonis Mt Gallego, Guadalcanal Island REPTILES OF THE SOLOMON ISLANDS

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81 Sphenomorphus tanneri (paratype) Mutahi, Bougainville Island

82 Sphenomorphus taylori (paratype) Bougainville Island COLOR PLATES 79/82

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83 Sphenomorphus transversus (holotype) Kunua, Bougainville Island

84 Sphenomorphus woodfordi Mboromole, Ngela Sule Island REPTILES OF THE SOLOMON ISLANDS

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85 Sphenomorphus sp. Malaupaina Island, Olu Malau Islands

86 Tribolonotus blanchardi Mboromole, Ngela Sule Island COLOR PLATES 83/86

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87 Tribolonotus blanchardi Mboromole, Ngela Sule Island

88 Tribolonotus ponceleti (hatchling) Offspring of parent lizards from Harapa, Shortland Island REPTILES OF THE SOLOMON ISLANDS

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89 Tribolonotus ponceleti (male) Harapa, Shortland Island

90 Tribolonotus ponceleti (female) Harapa, Shortland Island COLOR PLATES 87/90

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91 Tribolonotus pseudoponceleti Poroporo, Choiseul Island

92 Tribolonotus schmidti (male) Mt Austen, Guadalcanal Island REPTILES OF THE SOLOMON ISLANDS

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93 Tribolonotus schmidti (female) Mt Austen, Guadalcanal Island

94 Varanus indicus Malaupaina Island, Olu Malau Islands COLOR PLATES 91/94

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95 Varanus indicus (hatchling) Malaupaina Island, Olu Malau Islands

96 Varanus spinulosus San Jorge Island REPTILES OF THE SOLOMON ISLANDS

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97 Acutotyphlops infralabialis Mt Gallego, Guadalcanal Island

98 Ramphotyphlops angusticeps Malaupaina Island, Olu Malau Islands COLOR PLATES 95/98

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99 Ramphotyphlops becki Mt Austen, Guadalcanal Island

100 Ramphotyphlops braminus Mbua Valley, Guadalcanal Island REPTILES OF THE SOLOMON ISLANDS

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101 Ramphotyphlops depressus Teopasino, Bougainville Island

102 Ramphotyphlops depressus Mt Austen, Guadalcanal Island COLOR PLATES 99/102

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103 Candoia aspera New Ireland, Papua New Guinea

104 Candoia bibroni Malaupaina Island, Olu Malau Islands REPTILES OF THE SOLOMON ISLANDS

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105 Candoia bibroni Malaupaina Island, Olu Malau Islands

106 Candoia bibroni Lata, Santa Cruz Island COLOR PLATES 103/106

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107 Candoia bibroni Lomlom, Reef Islands

108 Candoia paulsoni Teopasino, Bougainville Island REPTILES OF THE SOLOMON ISLANDS

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109 Candoia paulsoni Kola’a Ridge, Guadalcanal Island

110 Candoia paulsoni Mamara, Guadalcanal Island COLOR PLATES 107/110

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111 Candoia paulsoni Marau, Guadalcanal Island

112 Candoia paulsoni Santa Catalina Island REPTILES OF THE SOLOMON ISLANDS

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113 Acrochordus granulatus Mboli Passage, Ngela Islands

114 Boiga irregularis Mbua Valley, Guadalcanal Island COLOR PLATES 111/114

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115 Boiga irregularis (hatchling) Tenaru, Guadalcanal Island

116 Dendrelaphis salomonis Mt Austen, Guadalcanal Island REPTILES OF THE SOLOMON ISLANDS

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117 Loveridgelaps elapoides Mboromole, Ngela Sule Island

118 Parapistocalamus hedigeri Bougainville Island COLOR PLATES 115/118

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119 Salomonelaps par Korovou, Shortland Island

120 Salomonelaps par Poroporo, Choiseul Island REPTILES OF THE SOLOMON ISLANDS

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121 Salomonelaps par Noro, New Georgia Island

122 Salomonelaps par Mt Austen, Guadalcanal Island COLOR PLATES 119/122

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123 Salomonelaps par (juvenile) Tenaru, Guadalcanal Island

124 Pelamis platurus Noumea, New Caledonia (photo: H. Cogger) REPTILES OF THE SOLOMON ISLANDS

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125 Laticauda colubrina Tulagi Island

126 Laticauda crockeri Lake Te-Nggano, Rennell Island COLOR PLATES 123/126

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127 Laticauda crockeri Lake Te-Nggano, Rennell Island

128 Laticauda laticaudata Vicinity of Heuru, Makira Island REPTILES OF THE SOLOMON ISLANDS

Hydrophis cyanocinctus (Daudin 1803) Distribution: This species ranges from the Persian Gulf, through the Indian Ocean and South China Sea, to the Indonesian archipelago and New Guinea. In the Solomons it is known from a single specimen collected off Guadalcanal (Brigham Young University no. 7861). McDowell and Cogger (pers. comms.) expressed doubt as to the correct identification of this specimen. On the basis of Tanner’s (1951) description, McDowell believed it to be referable to another species, possibly Hydrophis melanocephalus. However, Allen Allison of Bishop Museum arranged to have this specimen examined in 2001 by Harold Voris, who confirmed its identity as Hydrophis cyanocinctus. Description: The following is Tanner’s description of the specimen: “Rostral broader than deep with marginal grooves; nasal shorter than the frontal, twice as long as the suture between the prefrontals; prefrontals in contact with second supralabial; 1 preocular; 2 postoculars; temporals 3 and 1; 8 upper labials, second largest, third, fourth and fifth entering the eye; infralabials 10 and 9, both pair of chin shields in contact; body scale rows 27 anterior, 37 at midbody, 33 posterior near anus; anals 2 pairs; gastrosteges 334; urosteges 43; scales smooth and sub-imbricate. Colour of preserved specimen black above with 47 light bands extending from the dark back to the ventral surface. A single row of larger black gastrostege scales separates the light bands. The head and chin are light coloured. The tail for the length of 22 scales is black. Total length is 1028 (927+ 101)mm.” Natural History Notes: Probably similar to Hydrophis belcheri. Hydrophis ornatus (Gray, 1842) Distribution: Indian and western Pacific Oceans. In the Solomons it is known from specimens collected off Guadalcanal and Malaita. Description: Average total length around 1m. Body quite stout and not noticeably elongate. Head moderate. The tail is strongly compressed laterally. 1 preocular and 2 or 3 postoculars. The frontal is about as long as its distance from the snout. 7 or 8 supralabials, the third and fourth entering the eye. Midbody scales in 33-45 rows in males, and 39-55 in females. The body scales are subimbricate or juxtaposed. 209-260 ventrals in males, 236-312 in females, the ventrals about 2x as wide as the surrounding body scales. ELAPID SNAKES, Family ELAPIDAE

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Colour: Pale grey to dark grey above with a series of 30-60 dark transverse bars or blotches. A series of dark-edged ocellate markings is present laterally. The lower half of the body and the ventral area are yellowish to white. Natural History Notes: Probably similar to Hydrophis belcheri.

Genus Pelamis Daudin, 1803 A monotypic genus. Head shields enlarged, regular. Ventrals much reduced in size. Mental groove absent. Body scales small and juxtaposed. YELLOW-BELLIED SEA SNAKE Pelamis platurus (Linnaeus, 1766) Fig. 124 Distribution: Tropical and temperate regions of the Indian and Pacific Oceans. Description: Average total length 700mm, grows to over 1m. Head large and elongated. Body laterally compressed. Tail laterally compressed and paddle-shaped. No mental groove. 1 or 2 preoculars and 2 or 3 postoculars. Frontal large, as long as its distance from the snout. 7 to 8 supralabials, usually separated from the eye by subocular scales. The body is covered with small, more or less hexagonal scales. Midbody scales in 49 to 67 rows, these body scales are juxtaposed. 264 to 406 ventrals which are scarcely distinguishable from the surrounding body scales. Colour: The colour and pattern are variable. The commonest colour form is black above and yellow or cream below, these colours sharply differentiated midlaterally. The head is black above and the upper lip is usually yellow. The tail is usually mottled black and yellow. Natural History Notes: This species is not commonly seen in the Solomons, being an open sea dweller it rarely comes close to the coast, and the occasional specimens that are found on island shores are usually dead or dying as a result of illness or injury. Pelamis frequently follows driftlines; the edges of open sea currents where floating debris accumulates, feeding on the small fish (often larvae of pelagic species) that shelter under the debris. When approached on the surface I have seen Pelamis immediately dive vertically down into deep water. Solomon Islanders have told me that they have seen the brahminy kite Haliastur indus preying on Pelamis.

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Human fatalities have been attributed to the bite of Pelamis in other parts of the world. KEY TO THE SPECIES OF SOLOMON ISLANDS SEA KRAITS 1. Body distinctly banded, upper lip yellow, midbody scales in 21 or more rows, an azygous prefrontal almost always present: Laticauda colubrina – Banding on the body distinct, indistinct or absent, upper lip dark brown to black, midbody scales in 19-21 rows, no azygous prefrontal: 2 2. Body melanotic, banding varies from reasonably distinct to absent, midbody scales in 19-21 rows, Only found in Lake Te-Nggano, Rennell: Laticauda crockeri – Body not melanotic, distinctly banded, midbody scales in 19 rows, not found in Lake Te-Nggano, Rennell: Laticauda laticaudata

Genus Laticauda Laurenti, 1768 See description of the sea kraits in the family Elapidae above. YELLOW-LIPPED SEA KRAIT Laticauda colubrina (Schneider, 1799) Fig. 125 Distribution: Eastern Indian and western Pacific Oceans. Description: Average total length about 800mm, though in Lake TeNggano on Rennell Island these snakes can grow to almost 2m in length (personal observation). Head small, scarcely distinct from the neck. Body more or less round in cross section. Tail laterally compressed, paddleshaped. Nostril in a lateral position. An azygous prefrontal is almost always present between the prefrontals. 1 preocular, 2 postoculars. The frontal is much longer than its distance from the snout. 7 to 8 supralabials, the third and fourth entering the eye. Midbody scales in 21 to 25 rows. Ventrals well developed, 210 to 250. Anal divided. 37 to 47 subcaudals in males, 27 to 35 in females, all divided. Colour: Light or dark blue-grey above with 20 to 65 black bands of more or less uniform thickness or narrowing ventrally. Some or all of these ELAPID SNAKES, Family ELAPIDAE

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bands may be incomplete ventrally. The back of the head is black, usually continuous with a broad streak running from the posterior margin of the eye. The snout is yellow, this yellow colour extending back along the upper lip to the temporal region. Ventrally white, cream or light yellow. Natural History Notes: This common snake is an eel-specific feeder as an adult, though I have seen a juvenile specimen feeding diurnally on small fish (Gobidon sp.) that were sheltering in coral heads on a shallow reef. Adult snakes are usually nocturnal with an average foraging depth from 1 to 5m, though I have seen them at depths down to 20m. By day these snakes typically shelter on rocky islets under stones and in crevices. Juveniles are both diurnal and nocturnal and are sometimes seen foraging in tidal pools. On the island of Nusa Noke (which translates as Snake Island) in Vonavona Lagoon in the New Georgia Group, very large population densities of this species occur and on this island I have found these snakes diurnally under limestone slabs on the ground and in hollows in trees several metres above the ground. On Malaupaina Island in the Olu Malau group I once found a juvenile L. colubrina moving along a bush track during the day some 200m from the sea. Laticauda colubrina lays its eggs ashore, typically in well protected crevices in and under beach debris. In Lake Te-Nggano on Rennell Island, L. colubrina often congregates in large numbers in hollows in limestone rock on the small islets in the lake. It is an inoffensive snake, and although it is venomous it will normally make no attempt to bite, even when handled. In the Indo-Pacific a species of snake eel Myrichthys colubrina is often mistaken for Laticauda. This black and white banded eel mimics the snake in both colour pattern and behaviour. LAKE TE-NGGANO SEA KRAIT Laticauda crockeri Slevin, 1934 Figs. 126, 127 Distribution: Endemic to the Solomon Islands: Rennell. Type locality: Lake Te-Nggano, Rennell Island, Solomon Islands. Description: Average total length about 500mm for males and 650mm for females. Head small, scarcely distinct from the neck. Body more or less round in cross section. Tail laterally compressed, paddle-shaped. Nostril in a lateral position. No azygous prefontal. 1 preocular, 2 postoculars. The frontal is longer than its distance from the snout. 7 supralabials, the third and fourth

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entering the eye. Midbody scales in 19 rows (rarely 20 or 21 rows). Ventrals well developed, 193 to 210. Anal divided. 34 to 39 subcaudals. Colour: Brown to black dorsally, lighter brown laterally and ventrally. In most specimens lighter banding is distinguishable, at least laterally and ventrally. Natural History Notes: This dwarf melanotic sea krait is abundant in Lake Te-Nggano and forages through the limestone crevices in the shallow water around the periphery of the lake. It often collects in large numbers at the sites of freshwater springs that occur at several places around the edge of the lake. Although the lake is land-locked, the water is brackish. This snake very occasionally leaves the water and can be found moving amongst the short swampy grass at the lake edge. It is an inoffensive species and will make no attempt to bite, even when handled. The Rennellese people regard it as harmless. Laticauda crockeri apparently feeds exclusively on a species of small fish Eleotris fusca that is common in Lake Te-Nggano (personal underwater observation of wild specimens). The reproductive mode of this snake is presently unknown, though the Rennellese people claim it is a live-bearer. If this is correct, it would be the only non-oviparous Laticauda. BANDED SEA KRAIT Laticauda laticaudata (Linnaeus, 1758) Fig. 128 Distribution: Eastern Indian and western Pacific Oceans. To date this species is known in the Solomons from a single specimen which I found off the northern coast of Makira in 1975. Description: Average total length about 750mm, grows to about 1m. Head small, scarcely distinct from the neck. The body is more or less round in cross section. The tail is laterally compressed and paddle-shaped. Nostrils in a lateral position. No azygous prefrontal. 1 preocular and 2 postoculars. The frontal is longer than is distance from the snout. 7 supralabials, the third and fourth entering the eye. Midbody scales in 19 rows. Ventrals well developed, 225 to 245. Anal divided. 38 to 47 subcaudals in males, 30 to 35 in females, all divided. Colour: Dorsally light bluish grey with 25 to 70 black bands of more or less uniform thickness, some or all of these bands may be incomplete ventrally. ELAPID SNAKES, Family ELAPIDAE

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The head is posteriorly black and cream anteriorly. The area in front of the eyes and the labials are black. Ventrally white to cream. Natural History Notes: This species is apparently quite rare in the Solomons. The single known Solomons specimen (fig. 128) I found swimming on the surface at night about 100 metres offshore on the northern coast of Makira. Ineich and Laboute (2002) mention this species as being very similar to colubrina in its habits and feeding behaviour; although it shows somewhat less terrestrial activity than colubrina, and is less eel-specific, also feeding on gobies.

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LITERATURE CITED AND FURTHER REFERENCES ALLISON, A. 1997. Zoogeography of amphibians and reptiles of New Guinea and the Pacific Region, pp. 407–436 in A. Keast and S. E. Miller (eds.), The origin and evolution of Pacific Island biotas, New Guinea to Eastern Polynesia: Patterns and processes. SPB Academic Publishing, Amsterdam. BARBOUR, T. 1921. Reptiles and amphibians from the British Solomon Islands. Proc. New Eng. Zool. Club 7: 91–112. BÖHME, W., P. KAI and T. ZIEGLER. 2002. Another new member of the Varanus (Euprepiosaurus) indicus group (Sauria, Varanidae): an undescribed species from Rennell Island, Solomon Islands. Salamandra. 38 (1):15–26. BOULENGER, G.A. 1884. Diagnoses of new reptiles and batrachians from the Solomon Islands, collected and presented to the British Museum by H.B. Guppy, Esq., M.B., H.M.S. Lark. Proc. Zool. Soc. Lond. 210–213. BOULENGER, G.A. 1885-1887a. Catalogue of the lizards in the British Museum (N.H.), Vols. 1–3, British Museum (N.H.) London.Boulenger, G.A. 1887 b. Second contribution to the herpetology of the Solomon Islands. Proc. Zool. Soc. Lond. 333–338. BOULENGER, G.A. 1893–1896. Catalogue of the snakes in the British Museum (N.H.), Vols.1–3, British Museum (N.H.) London. BROWN, W.C. 1964. The status of Pseudogekko shebae and observations on the geckos of the Solomon Islands. Breviora 203: 1–4. BROWN, W.C. 1991. Lizards of the genus Emoia (Scincidae) with observations on their evolution and biogeography. Mem. Cal. Acad. Sci. 15: (1–94). BROWN, W.C. and M. MCCOY. 1980. A new species of gecko of the genus Cyrtodactylus from Guadalcanal Island, Solomon Islands. Herpetologica 36 (1): 66–69. LITERATURE CITED AND FURTHER REFERENCES

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BROWN, W.C., M. MCCOY and G. RODDA. 1992. A new Lepidodactylus (Reptilia: Gekkonidae) from Guadalcanal Island, Solomons. Proc. Biol. Soc. Wash. 105 (3): 440–442. BROWN, W.C., and F. PARKER. 1977. Lizards of the genus Lepidodactylus (Gekkonidae) from the Indo-Australian Archipelago and the islands of the Pacific, with descriptions of new species. Proc. Cal. Acad. Sci. (4th ser.) 41 (8): 253–265 BROWN, W.C. and V.M.TANNER. 1949. Rediscovery of the genus Pseudogekko with description of a new species from the Solomon Islands. Great Basin Nat. 9 (3–4): 41–45. BRUNA, E.M., R.N. FISHER and T.J. CASE. 1996. New evidence of habitat segregation between two cryptic species of Pacific skinks (Emoia cyanura and E. impar). Copeia 1996: 998–1005. BURT, C.E. 1930. Herpetological results of the Whitney South Sea expedition IV. Descriptions of new species of lizards from the Pacific Islands (Scincidae). Am. Mus. Nov. 427: 1–3. BURT, C.E. and M.D. BURT. 1932. Herpetological results of the Whitney South Sea expedition VI. Bull. Am. Mus. Nat. Hist. 63: 461–597. BUSTARD, R. 1970. Activity cycle of the tropical house gecko, Hemidactylus frenatus. Copeia 1970 (1): 173–176. COGGER, H.G. 1992. Reptiles and Amphibians of Australia 5th Edition. Reed Books, Sydney. COGGER, H., H. HEATWOLE, Y. ISHIKAWA, M. MCCOY, N. TAMIYA and T. TERUCHI. 1987. The status and natural history of the Rennell Island Sea Krait, Laticauda crockeri (Serpentes: Laticaudidae). J. Herp. 21 (4): 255–266. DE ROOIJ, N. 1915. The Reptiles of the Indo-Australian Archipelago. Vol.1. Lacertilia, Chelonia, Emydosauria. Leiden, E.J. Brill. DE ROOIJ, N. 1917. The Reptiles of the Indo-Australian Archipelago. Vol. 2. Ophidia. Leiden, E.J. Brill.

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FISHER, R. N. 1997. Dispersal and evolution of the Pacific basin gekkonid lizards Gehyra oceanica and Gehyra mutilata. Evolution 51 (3): 906–921. GREER, A.E. 1968. Clutch size in the scincid genus Emoia. Copeia 1968 (2): 417–418. GREER, A.E. 1970. The relationships of the skinks referred to the genus Dasia. Breviora 348: 1–30. GREER, A.E. 1974. The generic relationship of the scincid lizard genus Leiolopisma and its relatives. Aust. J. Zool. Suppl. 31: 1–67. GREER, A.E. and F. Parker. 1967a. A new scincid lizard from the northern Solomon Islands. Breviora 275: 1–21. GREER, A.E. and F. PARKER. 1967b. A second skink with fragmented head scales from Bougainville, Solomon Islands. Breviora 279: 1–12. GREER, A.E. and F. PARKER. 1968a. A new species of Tribolonotus (Lacertilia: Scincidae) from Bougainville and Buka, Solomon Islands, with comments on the biology of the genus. Breviora 291: 1–23. GREER, A.E. and F. PARKER. 1968b. Geomyersia glabra, a new genus and species of scincid lizard from Bougainville, Solomon Islands, with comments on the relationship of some lygosomine genera. Breviora 302: 1–17. GREER, A.E. and F. PARKER. 1971. A new scincid lizard from Bougainville, Solomon Islands. Breviora 364: 1–11. HARLOW, P. and R. SHINE 1992. Food habits and reproductive biology of the Pacific Island boas. (Candoia). J. Herp. 26 (1): 60–66. HEATWOLE, H. 1975. Biogeography of reptiles of some of the islands and cays of eastern Papua New Guinea. Atoll Research Bulletin 180. The Smithsonian Institution. INEICH, I. and P. LABOUTE 2002. Sea Snakes of New Caledonia. IRD Editions, Paris. INEICH, I. and G. ZUG. 1991. Nomenclatural status of Emoia cyanura (Lacertilia, Scincidae) populations in the central Pacific. Copeia 1991 (4): 1132–1136. LITERATURE CITED AND FURTHER REFERENCES

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KINGHORN, J.R. 1928. Herpetology of the Solomon Islands. Rec. Aust. Mus. 16 (3): 123–178. KINGHORN, J.R. 1937. A new species of skink from the Solomon Islands. Rec. Aust. Mus. 20 (1): 1–2. KLUGE, A.G. 1966. Phylogenetic relationships of the gekkonid lizard genera Lepidodactylus Fitzinger, Hemiphyllodactylus Bleeker and Pseudogekko Taylor. Phil. J. Sci. 95 (3): 331–352. KÖHLER, G. 1997. Eine neue Unterart des Wickelschwanzskinkes Corucia zebrata von Bougainville, Papua-Neuguinea. Salamandra. 33(1): 61–68. KRAUS, F. and A. ALLISON 2006. A new species of Cyrtodactylus (Lacertilia: Gekkonidae) from Papua New Guinea. Zootaxa 1247: 57-68. LEARY, T. 1991. A review of terrestrial wildlife trade originating from Solomon Islands. Aust. Zool. 27 (1 & 2): 20–27. LOVERIDGE, A. 1948. New Guinea reptiles and amphibians in the Museum of Comparative Zoology and United States National Museum. Bull. Mus. Comp. Zool. 101: 305–430. MANTHEY, U. and W. DENZER. 2006. A revision of the Melanesian-Australian angle-headed lizards of the genus Hypsilurus (Sauria: Agamidae: Amphibolurinae), with description of four new species and one new subspecies. Hamadryad 30 (1 & 2): 1–40. MCCOY, M. 1978. The agamid Gonocephalus godeffroyi in the Solomon Islands. Herpetofauna 10 (1): 2–4. MCCOY, M. 1980. Reptiles of the Solomon Islands. Wau Ecology Institute, Handbook No. 7 MCCOY, M. and P. WEBBER. 1984. Two new species of scincid lizards of the genus Emoia from Santa Cruz and Duff Islands, Solomon Islands. Copeia 1984: 571–578. MCDOWELL, S.B. 1969. On the status and relationship of the Solomon Islands elapid snakes. J. Zool. Lond. 191: 145–190.

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MCDOWELL, S.B. 1974. A catalogue of the snakes of New Guinea and the Solomons, with special reference to those in the Bernice P. Bishop Museum, Part 1. Scolecophidia. J. Herp. 8 (1): 1–-57. MCDOWELL, S.B. 1979. A catalogue of the snakes of New Guinea and the Solomons, with special reference to those in the Bernice P. Bishop Museum, Part 111. Boinae and Acrochordoidea (Reptilia, Serpentes). J. Herp. 13 (1): 1–92. MERTENS, R. 1931. Ablepharus boutonii (Desjardin) und seine geographische Variation. Zool. Jahrb. Abt. Syst. 61: 63–210. MERTENS, R. 1941. Die Familie der Warane (Varanidae). Abh. Senck. Nat. Ges. 462: 1–116. O’SHEA, M. 1996. A Guide to the Snakes of Papua New Guinea. Independent Publishing, Port Moresby, Papua New Guinea. O’SHEA, M. 2005. Venomous Snakes of the World. New Holland, London. PARKER, H.W. 1925. Notes on lizards from the South Pacific islands. Ann. Mag. Nat. Hist. (9) 15: 298–300. READ, J. L. and K. MOSEBY. 2006. Vertebrates of Tetepare Island, Solomon Islands. Pacific Science. 60 (1) 69–79. SCHMIDT, K.P. 1932. Reptiles and amphibians from the Solomon Islands. Pub. Field Mus. Nat. Hist. 18: 175–190. SCHMIDT, K.P. and C.E. BURT. 1930. Herpetological results of the Whitney South Sea Expedition V. Description of Emoia sanfordi, a new lizard from islands of the Western Pacific (Scincidae). Am. Mus. Nov. 436: 1–3. SHEA, G. and V. WALLACH. 2000. Reexamination of an anomalous distribution: resurrection of Ramphotyphlops becki (Serpentes: Typhlopidae). Pacific Science 54 (1): 70–74 SMITH, H.M., D. CHISZAR, K. TEPEDELEN and F. VAN BREUKELEN. 2001. A revision of the bevelnosed boas (Candoia carinata complex) (Reptilia: Serpentes). Hamadryad. 26 (2): 283–315. LITERATURE CITED AND FURTHER REFERENCES

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SLEVIN, J.R. 1934. The Templeton Crocker Expedition to western Polynesia and Melanesian islands 1933. No. 15. Notes on the reptiles and amphibians with the description of a new species of sea snake. Proc. Cal. Acad. Sci. (4th series) 21 (15): 183–188. SPRACKLAND, R.G. 1994. Rediscovery and taxonomic review of Varanus indicus spinulosus Mertens, 1941. Herpetofauna, 24 (2): 33–39. TANNER, V.M. 1951. Pacific Islands Herpetology No. V., Guadalcanal, Solomon Islands, a checklist of species. Great Basin Nat. 11 (3-4): 53–86. VOLSOE, H. 1956. Herpetology of Rennell Island, in The Natural History of Rennell Island, British Solomon Islands, Vol. I., Copenhagen. UETZ, P. 2006. The EMBL/EBI Reptile Database. (online: www.reptile-database.org) WAITE, E.R. 1905. Additions to the lacertilian fauna of the Solomon Islands. Rec. Aust. Mus. 6: 13–16. WALLACH, V. 1995. A new genus for the Ramphotyphlops subocularis species group (Serpentes: Typhlopidae), with description of a new species. Asiatic Herp. Res. (6): 132–150. WALLACH, V. 2003. Scolecophidia miscellanea. Hamadryad. 27(2): 222–240 WEBSTER, T.P. 1969. Aspects of the morphological and ecological variation in the cyanura group of the lizard genus Emoia (Sauria: Scincidae) in the Solomon Islands. Unpublished B.A. Honours thesis (Biology), Harvard University, Cambridge, Massachusetts. WILLIAMS, E.E. and F. Parker. 1964. The snake genus Parapistocalamus on Bougainville, Solomon Islands. Senk. Biol. 45: 543–552. ZWEIFEL, R.G. 1966. A new lizard of the genus Tribolonotus (Scincidae) from New Britain. Am. Mus. Nov. 2264: 1–12. ZUG, G. 2004. Systematics of the Carlia “fusca” lizards (Squamata: Scincidae) of New Guinea and nearby islands. Bishop Mus. Bull. Zool. 5: 1–84.

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ZUG, G.R., L.J. VITT and J.P. CALDWELL. (2001). Herpetology 2nd Edition Academic Press, San Diego, London ZUG, G. and B.R. MOON. 1995. Systematics of the Pacific slender-toed geckos Nactus pelagicus complex: Oceania, Vanuatu, and Solomon Islands populations. Herpetologica 51 (1): 77–90.

LITERATURE CITED AND FURTHER REFERENCES

125

GLOSSARY Abdominal glands: Glands, usually paired, lying towards the posterior venter of some species of skinks of the genus Tribolonotus. These glands are usually covered by enlarged scales. Adpress: To press flat against the body. When the limbs are adpressed, the forelimb is pressed backwards against the body and the hindlimb is pressed forwards against the body. Annuli: Rings or ring-like structures or patterns. Usually refers to scales. Anterior: Pertaining to the front, or head end of an animal. Arboreal: Dwelling in trees. Auricular lobules: Small protruding scales at the anterior edge of the ear opening in some skinks. Azygous: Not one of a pair, eg. an azygous scale. Bifid: Branching into two, forked. Bridge: In turtles, that part of the shell connecting the carapace to the plastron. Canthus rostralis: The ridge formed between the side and the top of the snout. Carapace: The top side of the shell in turtles. Carinate: Having a ridge or keel, usually refers to scales. Caudal: Pertaining to the tail. Circumnasals: In geckos, the enlarged scales (other than the rostral and labials) surrounding the nostril. Cloaca: The chamber into which the urinary, digestive and reproductive ducts open. Crepuscular: see Diurnal Cryptozoic: A secretive existence, living hidden under leaves, logs etc. Digital expansion: In geckos, that portion of the fingers and toes that is more or less distinctly dilated. Distal: That portion of a limb, digit, tail etc. that is furthest from its point of attachment to the body. Diurnal: Active during the day as opposed to nocturnal (active during the night) or crepuscular (active at dawn and/or dusk). Dorsal: Pertaining to the back (dorsum).

126

GLOSSARY

Dorsolateral: The area of the body where the side meets the back. Endemic (endemism): Occurring in a particular country or region and not elsewhere. Fossorial: Burrowing, an animal of burrowing habits. Front-fanged: In venomous snakes, having the fangs situated at the front of the maxillary bone. Gastrosteges: The ventral scales, the broad belly scales of snakes. Gular fold: A transverse fold of skin under the neck. Hemipenes: The paired male copulatory organs of snakes and lizards. Turtles and crocodiles have a single penis. Herbivorous: Feeding on plant material. Herpetology: The study of reptiles and amphibians. Imbricate: Overlapping one another, eg. imbricate scales. Juxtaposed: Lying side by side, not overlapping. Keel: A ridge, usually refers to keeled scales. Labial: 1. Pertaining to the lips. 2. The scales on the lips, supralabials are situated along the edge of the upper lip, infralabials along the lower lip. Lamellae: The ridges under the digits in lizards. Lateral: Pertaining to the sides of the body. Lateral fold: A longitudinal fold of skin along the lower edge of the side of the body. Littoral: Pertaining to the seashore. Mandible: The lower jaw. Maxillary: The upper jaw. Melanotic: Dark coloured as a result of an exceptional development in the amount of black pigment. Mental: 1. Pertaining to the chin. 2. The scale at the tip of the chin. Mental groove: A groove on the underside of the chin. Monogeneric: A family containing a single genus. Monotypic: A genus containing a single species. Montane: Pertaining to mountainous areas. Nasal cleft: A groove partly or completely dividing the nasal scales. Neurotoxic: A venom or poison affecting the nervous system. GLOSSARY

127

Nocturnal: see Diurnal Nuchal: 1. Pertaining to the neck. 2. The enlarged scales (usually paired) on the top of the neck. Occiput (occipital region): The area at the back of the head. Orbit: The cavity in the skull housing the eyeball. Oviparous: Egg-laying. Palpebral disc: A transparent disc on the lower eyelid of some skinks. Parietal eye: An internal structure situated in the parietal region of the skull and thought to play a role in thermoregulation in some lizards. Its presence is usually indicated externally by a small, darkly pigmented spot. Parthenogenesis: Reproduction without fertilisation of the ovum by a male. Pelagic: Dwelling in the open sea. Pentadactyl: Having five digits on a limb. Phalanx (pl. phalanges): The individual bones of a digit. Plastron: The underside of the shell in turtles. Pores: The opening of glands that are situated on different parts of the body of some lizards. Femoral pores are situated across the insides of the thighs, precloacal pores anterior to the vent. Palmar pores are situated on the insides of the hands, plantar pores on the inside of the feet. Posterior: Pertaining to the rear, or tail end of an animal. Postmental: The scale(s) situated immediately posterior to the mental scale. Precloacals: In lizards, the scales situated immediately anterior to the vent. Preocular: The scale(s) immediately in front of the eye. Proximal: That portion of a limb, digit, tail etc. that is closest to its point of attachment to the body. Rear-fanged: In venomous snakes, having the fangs situated at the rear of the maxillary bone. Rostral: The scale at the tip of the upper jaw. Sexual dimorphism: A difference between male and female of the same species. Usually a difference in size or proportion or colour (sexual dichromatism). Subcaudal: 1. The underside of the tail. 2. The scales (usually enlarged) on the underside of the tail. Supraciliaries: The small scales at the edge of the upper eyelid. Supranasal: A small scale situated immediately above the nasal scale.

128

GLOSSARY

SVL: Snout-vent length, the standard measurement of lizards. The distance from the snout to the vent measured ventrally. Terrestrial: Dwelling on the ground. Tubercle: A small protuberance, eg. a small conical scale. Tympanum: The resonating membrane of the ear. Urosteges: The enlarged scales under the tail (subcaudals). Vent: The external opening of the cloaca. Venter: The belly or underside of the body. Ventral: 1. Pertaining to the venter. 2. The broad belly scales of snakes. Vertebral: 1. That part of the body lying over the backbone. 2. In turtles, the dermal plates down the centre of the carapace. Viviparous: giving birth to young; live-bearing.

GLOSSARY

129

ISLAND CHECK LISTS BOUGAINVILLE Agamidae Hypsilurus longii Gekkonidae Cyrtodactylus louisiadensis Gehyra oceanica Gekko vittatus Hemidactylus frenatus Hemiphyllodactylus typus Lepidodactylus guppyi Lepidodactylus lugubris Lepidodactylus mutahi Nactus multicarinatus Scincidae Carlia mysi Corucia zebrata Emoia atrocostata Emoia caeruleocauda Emoia cyanogaster Emoia cyanura Emoia flavigularis Emoia impar Emoia jakati Emoia nigra Emoia pseudocyanura Eugongylus albofasciolatus Geomyersia glabra Lamprolepis smaragdina Lipinia noctua Prasinohaema virens Sphenomorphus concinnatus

130

ISLAND CHECK LISTS

Sphenomorphus cranei Sphenomorphus fragosus Sphenomorphus solomonis Sphenomorphus tanneri Sphenomorphus taylori Sphenomorphus transversus Sphenomorphus woodfordi Tribolonotus blanchardi Tribolonotus ponceleti (?) Tribolonotus pseudoponceleti Varanidae Varanus indicus Typhlopidae Acutotyphlops infralabialis Acutotyphlops kunuaensis Acutotyphlops solomonis Ramphotyphlops braminus Ramphotyphlops depressus Ramphotyphlops mansuetus Boidae Candoia aspera (?) Candoia paulsoni Acrochordidae Acrochordus granulatus Colubridae Boiga irregularis Dendrelaphis salomonis Elapidae Parapistocalamus hedigeri SHORTLAND ISLANDS (Shortland, Fauro, Mono) Agamidae Hypsilurus longii (?) Gekkonidae Cyrtodactylus louisiadensis Gehyra oceanica Gekko vittatus ISLAND CHECK LISTS

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Hemidactylus frenatus Lepidodactylus guppyi Lepidodactylus lugubris Nactus multicarinatus Scincidae Carlia mysi Corucia zebrata Emoia atrocostata Emoia caeruleocauda Emoia cyanogaster Emoia cyanura Emoia flavigularis Emoia impar Emoia nigra Emoia pseudocyanura Eugongylus albofasciolatus Eugongylus rufescens Geomyersia glabra Lamprolepis smaragdina Lipinia noctua Prasinohaema virens Sphenomorphus concinnatus Sphenomorphus cranei Sphenomorphus solomonis Sphenomorphus tanneri Sphenomorphus woodfordi Tribolonotus ponceleti Varanidae Varanus indicus Typhlopidae Ramphotyphlops depressus Boidae Candoia paulsoni Acrochordidae Acrochordus granulatus Colubridae Boiga irregularis

132

ISLAND CHECK LISTS

Dendrelaphis salomonis Elapidae Loveridgelaps elapoides Salomonelaps par CHOISEUL Gekkonidae Gehyra oceanica Hemidactylus frenatus Lepidodactylus lugubris Nactus multicarinatus Scincidae Corucia zebrata Emoia atrocostata Emoia caeruleocauda Emoia cyanogaster Emoia cyanura Emoia flavigularis Emoia nigra Emoia pseudocyanura Eugongylus albofasciolatus Lamprolepis smaragdina Sphenomorphus concinnatus Sphenomorphus solomonis Sphenomorphus tanneri Tribolonotus blanchardi Tribolonotus pseudoponceleti Varanidae Varanus indicus Boidae Candoia paulsoni Colubridae Boiga irregularis Dendrelaphis salomonis Elapidae Loveridgelaps elapoides Salomonelaps par ISLAND CHECK LISTS

133

VELLA LAVELLA Gekkonidae Gekko vittatus Lepidodactylus lugubris Nactus multicarinatus Scincidae Corucia zebrata Emoia atrocostata Emoia caeruleocauda Emoia cyanogaster Emoia cyanura Emoia nigra Eugongylus albofasciolatus Lamprolepis smaragdina Prasinohaema virens Sphenomorphus concinnatus Sphenomorphus cranei Varanidae Varanus indicus Typhlopidae Ramphotyphlops depressus Boidae Candoia paulsoni Colubridae Boiga irregularis Dendrelaphis salomonis Elapidae Loveridgelaps elapoides Salomonelaps par GIZO Gekkonidae Gekko vittatus Hemidactylus frenatus Nactus multicarinatus Scincidae Emoia atrocostata

134

ISLAND CHECK LISTS

Emoia cyanura Emoia schmidti Lamprolepis smaragdina Sphenomorphus concinnatus Varanidae Varanus indicus Boidae Candoia paulsoni Colubridae Dendrelaphis salomonis Elapidae Loveridgelaps elapoides Salomonelaps par RANONGGA Gekkonidae Lepidodactylus lugubris Nactus multicarinatus Scincidae Emoia atrocostata Emoia caeruleocauda Emoia cyanogaster Emoia cyanura Emoia nigra Lamprolepis smaragdina Sphenomorphus concinnatus Typhlopidae Ramphotyphlops depressus Boidae Candoia paulsoni Colubridae Boiga irregularis Dendrelaphis salomonis Elapidae Salomonelaps par

ISLAND CHECK LISTS

135

KOLOMBANGARA Gekkonidae Lepidodactylus guppyi Scincidae Emoia caeruleocauda Emoia schmidti Lamprolepis smaragdina Sphenomorphus bignelli Sphenomorphus concinnatus Varanidae Varanus indicus Typhlopidae Ramphotyphlops depressus Boidae Candoia paulsoni Colubridae Dendrelaphis salomonis Elapidae Salomonelaps par NEW GEORGIA Gekkonidae Cyrtodactylus louisiadensis Gehyra oceanica Gekko vittatus Hemidactylus frenatus Lepidodactylus guppyi Lepidodactylus lugubris Nactus multicarinatus Scincidae Corucia zebrata Emoia atrocostata Emoia caeruleocauda Emoia cyanogaster Emoia nigra Emoia schmidti Eugongylus albofasciolatus

136

ISLAND CHECK LISTS

Lamprolepis smaragdina Lipinia noctua Prasinohaema virens Sphenomorphus cranei Sphenomorphus solomonis Varanidae Varanus indicus Typhlopidae Acutotyphlops infralabialis Ramphotyphlops depressus Boidae Candoia paulsoni Acrochordidae Acrochordus granulatus Colubridae Dendrelaphis salomonis Elapidae Salomonelaps par VANGUNU Gekkonidae Gehyra oceanica Gekko vittatus Nactus multicarinatus Scincidae Corucia zebrata Emoia atrocostata Emoia caeruleocauda Emoia cyanogaster Emoia cyanura Emoia nigra Emoia schmidti Eugongylus albofasciolatus Lamprolepis smaragdina Lipinia noctua Prasinohaema virens Sphenomorphus bignelli ISLAND CHECK LISTS

137

Sphenomorphus concinnatus Sphenomorphus cranei Varanidae Varanus indicus Boidae Candoia paulsoni Colubridae Boiga irregularis Dendrelaphis salomonis Elapidae Salomonelaps par ISABEL Agamidae Hypsilurus longii Gekkonidae Gehyra oceanica Hemidactylus frenatus Lepidodactylus guppyi Nactus multicarinatus Scincidae Corucia zebrata Emoia atrocostata Emoia cyanogaster Emoia flavigularis Eugongylus albofasciolatus Lamprolepis smaragdina Sphenomorphus concinnatus Sphenomorphus cranei Sphenomorphus solomonis Tribolonotus ponceleti Varanidae Varanus indicus Varanus spinulosus Typhlopidae Ramphotyphlops depressus

138

ISLAND CHECK LISTS

Boidae Candoia paulsoni Acrochordidae Acrochordus granulatus Colubridae Boiga irregularis Dendrelaphis salomonis Elapidae Loveridgelaps elapoides Salomonelaps par GUADALCANAL Gekkonidae Cyrtodactylus biordinis Cyrtodactylus louisiadensis Gehyra mutilata Gehyra oceanica Gekko vittatus Hemidactylus frenatus Hemiphyllodactylus typus Lepidodactylus flaviocularis Lepidodactylus guppyi Lepidodactylus lugubris Lepidodactylus shebae Nactus multicarinatus Scincidae Corucia zebrata Emoia atrocostata Emoia cyanogaster Emoia cyanura Emoia nigra Emoia pseudocyanura Eugongylus albofasciolatus Lamprolepis smaragdina Lipinia noctua Prasinohaema virens “Sphenomorphus” bignelli ISLAND CHECK LISTS

139

Sphenomorphus concinnatus Sphenomorphus cranei Sphenomorphus solomonis Tribolonotus schmidti Varanidae Varanus indicus Typhlopidae Acutotyphlops infralabialis Ramphotyphlops angusticeps Ramphotyphlops becki Ramphotyphlops braminus Ramphotyphlops depressus Boidae Candoia paulsoni Acrochordidae Acrochordus granulatus Colubridae Boiga irregularis Dendrelaphis salomonis Elapidae Loveridgelaps elapoides Salomonelaps par NGELA Gekkonidae Gehyra oceanica Gekko vittatus Hemidactylus frenatus Lepidodactylus guppyi Lepidodactylus lugubris Nactus multicarinatus Scincidae Corucia zebrata Emoia atrocostata Emoia cyanogaster Emoia cyanura Emoia flavigularis

140

ISLAND CHECK LISTS

Emoia nigra Emoia pseudocyanura Eugongylus albofasciolatus Geomyersia glabra Lipinia noctua Prasinohaema virens “Sphenomorphus” bignelli Sphenomorphus concinnatus Sphenomorphus cranei Sphenomorphus solomonis Sphenomorphus woodfordi Tribolonotus blanchardi Varanidae Varanus indicus Typhlopidae Acutotyphlops infralabialis Ramphotyphlops depressus Boidae Candoia paulsoni Acrochordidae Acrochordus granulatus Colubridae Boiga irregularis Dendrelaphis salomonis Elapidae Loveridgelaps elapoides Salomonelaps par RENNELL Gekkonidae Gehyra oceanica Gekko vittatus Lepidodactylus lugubris Nactus multicarinatus Scincidae Cryptoblepharus poecilopleurus Emoia atrocostata ISLAND CHECK LISTS

141

Emoia cyanogaster Emoia rennellensis Eugongylus rufescens Varanidae Varanus indicus Typhlopidae Ramphotyphlops angusticeps Boidae Candoia bibroni Candoia paulsoni Elapidae Laticauda colubrina (in Lake Te-Nggano) Laticauda crockeri (in Lake Te-Nggano) MALAITA Gekkonidae Cyrtodactylus louisiadensis Gehyra oceanica Hemidactylus frenatus Lepidodactylus guppyi Lepidodactylus lugubris Nactus multicarinatus Scincidae Corucia zebrata Emoia atrocostata Emoia cyanogaster Emoia cyanura Emoia flavigularis Emoia nigra Emoia pseudocyanura Eugongylus albofasciolatus Lamprolepis smaragdina Prasinohaema virens Sphenomorphus bignelli Sphenomorphus concinnatus Sphenomorphus solomonis

142

ISLAND CHECK LISTS

Varanidae Varanus indicus Typhlopidae Acutotyphlops infralabialis Ramphotyphlops angusticeps Ramphotyphlops depressus Boidae Candoia paulsoni Acrochordidae Acrochordus granulatus Colubridae Boiga irregularis Dendrelaphis salomonis Elapidae Loveridgelaps elapoides Salomonelaps par MAKIRA Agamidae Hypsilurus macrolepis Gekkonidae Gehyra mutilata Gekko vittatus Lepidodactylus guppyi Lepidodactylus lugubris Nactus multicarinatus Scincidae Corucia zebrata Emoia atrocostata Emoia caeruleocauda Emoia cyanogaster Emoia cyanura Emoia maculata Emoia nigra Eugongylus albofasciolatus Lipinia noctua Prasinohaema virens ISLAND CHECK LISTS

143

Sphenomorphus woodfordi Sphenomorphus sp. Varanidae Varanus indicus Typhlopidae Ramphotyphlops mansuetus Ramphotyphlops angusticeps Ramphotyphlops depressus Boidae Candoia bibroni Candoia paulsoni Colubridae Dendrelaphis salomonis OLU MALAU Agamidae Hypsilurus macrolepis Gekkonidae Gehyra oceanica Gekko vittatus Lepidodactylus guppyi Lepidodactylus lugubris Nactus multicarinatus Scincidae Emoia atrocostata Emoia cyanogaster Emoia cyanura Emoia maculata Emoia nigra Eugongylus albofasciolatus Lipinia noctua Prasinohaema virens Sphenomorphus sp. Varanidae Varanus indicus Typhlopidae Ramphotyphlops angusticeps

144

ISLAND CHECK LISTS

Ramphotyphlops depressus Boidae Candoia bibroni Candoia paulsoni Colubridae Dendrelaphis salomonis NENDO (SANTA CRUZ) Gekkonidae Gehyra mutilata Gehyra oceanica Gekko vittatus Hemidactylus frenatus Lepidodactylus guppyi Lepidodactylus lugubris Nactus multicarinatus Scincidae Emoia atrocostata Emoia caeruleocauda Emoia cyanogaster Emoia nigra Emoia rufilabialis Eugongylus albofasciolatus Lamprolepis smaragdina Prasinohaema virens Sphenomorphus solomonis Boidae Candoia bibroni Candoia paulsoni Colubridae Boiga irregularis Dendrelaphis salomonis

ISLAND CHECK LISTS

145

INDEX OF SCIENTIFIC NAMES Acrochordus granulatus 104 Acutotyphlops infralabialis 95 Acutotyphlops kunuaensis 95 Acutotyphlops solomonis 96 Boiga irregularis 105 Candoia aspera 101 Candoia bibroni 101 Candoia paulsoni 102 Caretta caretta 18 Carlia mysi 47 Chelonia mydas 19 Corucia zebrata 48 Cryptoblepharus poecilopleurus 50 Cyrtodactylus biordinis 29 Cyrtodactylus louisiadensis 30

Emoia rufilabialis 66 Emoia sanfordi 66 Emoia schmidti 67 Emoia taumakoensis 68 Eretmochelys imbricata 20 Eugongylus albofasciolatus 69 Eugongylus rufescens 70 Gehyra mutilata 31 Gehyra oceanica 32 Gekko vittatus 34 Geomyersia glabra 71

Dendrelaphis salomonis 106 Dermochelys coriacea 22

Hemidactylus frenatus 35 Hemiphyllodactylus typus 37 Hydrophis belcheri 112 Hydrophis cyanocinctus 113 Hydrophis ornatus 113 Hypsilurus longii 24 Hypsilurus macrolepis 25

Emoia atrocostata 54 Emoia caeruleocauda 55 Emoia cyanogaster 56 Emoia cyanura 57 Emoia flavigularis 58 Emoia impar 59 Emoia isolata 60 Emoia jakati 61 Emoia maculata 62 Emoia nigra 63 Emoia pseudocyanura 64 Emoia rennellensis 65

Lamprolepis smaragdina 72 Laticauda colubrina 115 Laticauda crockeri 116 Laticauda laticaudata 117 Lepidochelys olivacea 21 Lepidodactylus flaviocularis 38 Lepidodactylus guppyi 39 Lepidodactylus lugubris 40 Lepidodactylus mutahi 41 Lepidodactylus shebae 42 Lepidodactylus woodfordi 43 Lipinia noctua 74

146

INDEX OF SCIENTIFIC NAMES

Loveridgelaps elapoides 108 Nactus multicarinatus 43 Parapistocalamus hedigeri 110 Pelamis platurus 114 Prasinohaema virens 75 Ramphotyphlops angusticeps 96 Ramphotyphlops becki 97 Ramphotyphlops braminus 98 Ramphotyphlops depressus 99 Ramphotyphlops mansuetus 99 Salomonelaps par 110 Sphenomorphus bignelli 78 Sphenomorphus concinnatus 79

Sphenomorphus cranei 80 Sphenomorphus fragosus 80 Sphenomorphus solomonis 81 Sphenomorphus sp. 84 Sphenomorphus tanneri 82 Sphenomorphus taylori 82 Sphenomorphus transversus 83 Sphenomorphus woodfordi 83 Tribolonotus blanchardi 85 Tribolonotus ponceleti 87 Tribolonotus pseudoponceleti 88 Tribolonotus schmidti 89 Varanus indicus 91 Varanus spinulosus 92

INDEX OF SCIENTIFIC NAMES

147

148