Field Guide to Frogs of Western Australia [1 ed.] 9781920843915, 9781920843403

Citation preview

Western Australian Museum

Field Guide to

FROGS of w e s t e r n aus t r a l i a Fourth edition

M.J. Tyler and P. Doughty

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Field Guide to

FROGS o f W E S T E RN A u s t r a l i a Fourth edition

Litoria moorei, Emu Point (G. Harold)

BY MICHAEL J. TYLER & PAUL DOUGHTY

*29(510(172) :(67(51$8675$/,$

DEDICATION

John A. Moore (1915–2002) For his pioneering studies of Australian frogs in the 1950s (Moore 1961) and advocate for ‘science as a way of knowing’ (Moore 1993, 2002).

CONTENTS Preface. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 08 How The Book Works. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 09 The Western Australian frog fauna . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 Basic frog biology. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12 Size, form and features. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12 Eggs and spawn. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19 Tadpoles and their morphology. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20 Legislation applying to frogs in Western Australia. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21 Regional comparisons . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22 A regional approach to identification. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23 Accounts of fauna . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23 ACCOUNTS OF FROG GENERA AND SPECIES Arenophryne Family Myobatrachidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25 A. rotunda Northern Sandhill Frog . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26 A. xiphorhyncha Southern Sandhill Frog. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27 Crinia Family Myobatrachidae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29 C. bilingua Bilingual Froglet . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30 C. deserticola Chirping Froglet. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31 C. fimbriata Kimberley Froglet. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32 C. georgiana Quacking Frog. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33 C. glauerti Rattling or Clicking Froglet. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35 C. insignifera Squelching Froglet . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36 C. pseudinsignifera Bleating Froglet . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37 C. subinsignifera South Coast Froglet. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38 Cyclorana Family Hylidae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39 C. australis Giant Frog. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40 C. cryptotis Hidden-ear Frog. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41 C. cultripes Knife-footed Frog . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 42 C. longipes Long-footed Frog . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43 C. maini Sheep or Main’s Frog. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 44 C. platycephala Water-holding Frog . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45 C. vagitus Wailing Frog . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 46 Geocrinia Family Myobatrachidae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 47 G. alba White-bellied Frog. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 48 G. leai Ticking Frog. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 49 G. lutea Walpole Frog. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 50 G. rosea Roseate Frog. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51 G. vitellina Orange or Yellow-bellied Frog. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 52 Heleioporus Family Limnodynastidae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 53 H. albopunctatus Western Spotted Frog. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 54 H. barycragus Hooting Frog. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 55 H. eyrei Moaning Frog. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 56 H. inornatus Whooping Frog. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 57 H. psammophilus Sand Frog. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 58

Limnodynastes Family Limnodynastidae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 61 L. convexiusculus Marbled Frog . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 62 L. depressus Flat-headed Frog . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 63 L. dorsalis Western Banjo Frog. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 64 L. lignarius Woodworker or Carpenter Frog. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 65 Litoria Family Hylidae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 67 L. adelaidensis Slender Tree Frog . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 68 L. bicolor Northern Dwarf Tree Frog. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69 L. caerulea Green Tree Frog. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 70 L. cavernicola Cave Frog. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 71 L. coplandi Rock Frog . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 72 L. cyclorhyncha Spotted-thighed Frog. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 73 L. dahlii Aquatic Frog . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 74 L. inermis Bumpy Rocket Frog. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 75 L. meiriana Rockhole Frog. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 76 L. microbelos Javelin Frog. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 77 L. moorei Motorbike Frog. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 78 L. nasuta Striped Rocket Frog . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 79 L. pallida Pale Rocket Frog . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 80 L. rothii Northern Laughing Tree Frog . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 81 L. rubella Little Red Tree Frog. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 82 L. splendida Magnificent or Splendid Tree Frog. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 83 L. staccato Chattering Rock Frog. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 84 L. tornieri Black-shinned Rocket Frog. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 85 L. watjulumensis Wotjulum Frog. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 86 Metacrinia Family Myobatrachidae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 88 M. nichollsi Forest Toadlet. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 89 Myobatrachus Family Myobatrachidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 90 M. gouldii Turtle Frog . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 91 Neobatrachus Family Limnodynastidae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 93 N. albipes White-footed Trilling Frog. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 94 N. aquilonius Northern Burrowing Frog . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 95 N. centralis Desert Trilling Frog. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 96 N. fulvus Tawny Trilling Frog . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 97 N. kunapalari Wheatbelt Frog. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 98 N. pelobatoides Humming Frog . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 99 N. sutor Shoemaker Frog . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 100 N. wilsmorei Plonking Frog . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101 Notaden Family Limnodynastidae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 103 N. melanoscaphus Northern Spadefoot. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 104 N. nichollsi Desert Spadefoot. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 105 N. weigeli Kimberley Spadefoot. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 106 Platyplectrum Family Limnodynastidae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 107 P. ornatum Ornate Burrowing Frog. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 108 P. spenceri Centralian Burrowing Frog. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 109 Pseudophryne Family Myobatrachidae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 111 P. douglasi Gorge Toadlet. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 112

P. guentheri Crawling Toadlet . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 113 P. occidentalis Western Toadlet. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 114 Spicospina Family Myobatrachidae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 116 S. flammocaerulea Sunset Frog. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 117 Uperoleia Family Myobatrachidae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 119 U. aspera Derby Toadlet. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 120 U. borealis Northern Toadlet. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 121 U. crassa Fat Toadlet. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 122 U. glandulosa Glandular Toadlet . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 123 U. lithomoda Stonemason Toadlet . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 124 U. marmorata Marbled Toadlet. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 125 U. micra Tiny Toadlet . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 126 U. micromeles Tanami Toadlet. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 127 U. minima Small Toadlet. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 128 U. mjobergii West Kimberley Toadlet. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 129 U. russelli Northwest Toadlet. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 130 U. talpa Mole Toadlet. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 131 U. trachyderma Blacksoil Toadlet. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 132 Bufo Family Bufonidae (naturalised exotic). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 134 B. marinus Cane Toad. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 135 CHECKLISTS OF FROGS FOR URBAN CENTRES Southwest region

Geraldton. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 136 Perth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 136 Bunbury. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 136 Albany. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 136 Esperance. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 136 Arid zone



Port Hedland. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 136 Exmouth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 137 Newman . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 137 Kalgoorlie. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 137 Kimberley region



Kununurra. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 137 Derby. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 137 Broome. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 137

Glossary. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 138 Approaches to identification . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 141 Southwest region key. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 141 Arid zone key. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 144 Kimberley region key. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 146 References. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 150 Acknowledgements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 156 Author’s Notes. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 157 Index. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 158

Preface to FOURTH Edition What is it about frogs that makes them so intriguing to people? Is it their large eyes, adapted so they can see in the night? Is it their wonderful colours and markings, adapted so that birds won’t see them in the day? Perhaps it is their current conservation plight – threatened by human expansion, including introduced diseases and toads – coupled with their non-aggressive, even contemplative, nature that endears them to children and adults alike. Maybe it is just the hop – both surprising and comical at once – that does it. Whatever the reason, frogs enjoy a massive appeal with people and have become a part of our culture. Western Australia occupies the entire western third of the Austalian continent. The environments the state embraces include the ancient swamps and old-growth forests of the southwest region with its Mediterranean climate, the parallel red sand dunes of the arid zone that stretch as far as the eye can see, the cool quiet gorges of the rugged Pilbara region and the untamed Kimberley that comes alive in summer with monsoonal rains, flowing creeks and waterfalls. This is the stage for the stars of this field guide – the frogs of Western Australia. There are many highlights when discussing encounters with Western Australian frogs. Many Western Australian’s experience will be an encounter with a southwestern Motorbike Frog (Litoria moorei) – calling (‘accelerating’) away on a spring night from a neighbor’s pond, or perhaps looking for a way out of the laundry. A traveler might get a start from seeing so many Little Red Tree Frogs (L. rubella) in a Karijini shower block, or perhaps a large Magnificent Tree Frog (L. splendida) in a Kununurra caravan park. The acrobatic antics of Rockhole Frogs (L. meiriana) might be serendipitiously enjoyed by a Kimberley waterfall. For the more intrepid nature-lover or biologist in the bush, some of the joys of frogging might be getting ‘oozed’ by a Notaden species, being perplexed by the true identity of a Uperoleia encountered during a survey, inadvertently waking up a ‘cocooned’ burrowing frog while excavating a fire pit or being serenaded by a raucous chorus of Wotjulum Frogs (L. watjulumensis) well into the morning light. Yet one does not have to travel far to experience some spectacles of nature of the frog kind. For example, just out of Perth all five Western Australian species of the large burrowing frogs of the genus Heleioporus can be encountered in autumn – a situation of which eastern Australian and overseas froggers would be extremely jealous! On the way to ‘listening-out’ for Heleioporus species, the Perth frogger could also encounter Crawling Frogs (Pseudophyrne guentheri) and Ticking Frogs (Geocrinia leai), and later in winter come across choruses of hundreds of Quacking Frogs (Crinia georgiana) or Rattling Frogs (C. glauerti). In recent years, the idea of ‘frogs’ in the popular imagination has been tightly linked to ‘conservation’ and ‘extinction’. For a frog biologist, there is nothing sadder than listening to a recording of a call of an extinct species. Unfortunately, most news articles we read of are ‘bad news’ stories about frogs, especially impacts due to land clearing for developments, the spraying of toxic chemicals harmful to frogs to control weeds and insects, ‘disappearing’ species in eastern Australia owing to the exotic chytrid fungus and the imminent invasion of the Kimberley by the notorious South American Cane Toad (Bufo marinus). Despite the realilty of these scenarios, to stop there would be to miss the big picture – the wonderful, diverse and healthy status of nearly all species of frogs that can still be experienced today in the Western Australian bush.

8

How the book works This field guide was written primarily to provide the means necessary to identify frogs in the field. Identification of species is possible by using the keys to Western Australia’s three main regions at the back of the book, then referring to the descriptions, photographs and maps to arrive at the correct identification. Extra information is provided for a better understanding of a species’ habits and life-history, and a list of references for further reading (although we still know very little of many species). An introductory section on the biology of frogs is provided to give the reader a grasp of the morphological characters referred to in the text. There is a glossary of scientific terms at the end of the book. In the accounts of species, the frogs are presented in alphabetical order, with genus first, followed by species. We hope your encounters with frogs in the bush are enhanced by successful identification and through learning some of their habits through use of this field guide. Most importantly, it is the frogs themselves that are the true ‘book’ that is worth reading by the curious naturalist. Happy frogging!

9

THE WESTERN AUSTRALIAN FROG FAUNA In 1961 John Moore (a visiting Fulbright Scholar from the USA) published a monograph with a list of the frogs of Australia that included 94 species. Nearly 50 years later, this total has risen to 228. As we learn more information about Australian frogs, further species are discovered and formally described. The real diversity of the fauna is unlikely to be known for many years, and the total number of Australian frogs may well be over 250 species. The Western Australian frog fauna is comprised of three native families, with a fourth exotic family that has only recently invaded Western Australia. Hylidae (82 Australian species; 26 WA species) This is one of the largest families of amphibians with almost 900 species. Although members of this family are described collectively as ‘tree frogs’, they include a number of species that are terrestrial and are incapable of climbing. In Australia, species in the genus Cyclorana are adept at burrowing beneath the surface of the ground. Australian species are placed in the subfamily Pelodryadinae; the other two subfamilies (Hylinae and Phyllomedusinae) are also speciose, especially in the New World tropics (see Plötner et al. 2007). In Australia, hylids are represented by about 100 species placed in three genera: the large and diverse genus Litoria, the burrowing species of the genus Cyclorana and, in Queensland, a single species of Nyctimystes, a genus more widely represented in New Guinea. This taxonomy will undergo some major revisions in the coming years as the many ‘species-groups’ within Litoria are likely to be given generic status. Limnodynastidae (41 Australian species; 22 WA species) This is a group of large-bodied Gondwanan frogs that are widespread throughout Australia and parts of New Guinea. This family has traditionally been regarded as a subfamily within the Myobatrachidae, but recent work has supported its distinctiveness (e.g. Frost et al. 2006). Many species are excellent burrowers, but some tropical species are adept at climbing rocks (Limnodynastes lignarius, Notaden weigeli). It contains Limnodynastes, Adelotus, Heleioporus, Lechriodus, Neobatrachus, Notaden, Philoria and Platyplectrum. Except for Neobatrachus and Notaden, species in all genera produce foamy egg masses (although not all species do; e.g. Philoria loveridgei). Myobatrachidae (83 Australian species; 33 WA species) The Myobatrachidae is a group of generally small-bodied ground-dwelling or burrowing species, but includes a few small climbers as well. There are three especially large genera – Crinia (16 species), Pseudophryne (13) and Uperoleia (26). The other genera are Myobatrachus, Arenophryne, Assa, Geocrinia, Metacrinia, Paracrinia, Spicospina and Taudactylus and also includes the more distantly-related eastern genera Rheobatrachus and Mixophyes. Myobatrachids show a wide diversity of breeding modes, including gastric brooding, ‘marsupial’-like parental care, direct-developing eggs, terrestrial nests with non-feeding tadpoles, eggs laid in vegetation above water bodies, in addition to the typical aquatic eggs and tadpoles. Like the Limnodynastidae, they only occur in Australia and parts of southern New Guinea. Bufonidae (1 species in Australia – naturalized exotic) The genus represented by this family in Australia is native to South, Central and the extreme south of North America. In 1935, however, the Cane Toad (Bufo marinus) was introduced to Queensland from introduced Hawaiian populations to control native beetles that were attacking sugar cane (an introduced crop itself). The toad had no effect on the beetles, and has now spread throughout most of Queensland, northern New South Wales and the northern portion of the Northern Territory. At this writing Cane Toads have just crossed the Western Australian border, and is therefore worth including in this field guide to assist in identification. 10

THE WESTERN AUSTRALIAN FROG FAUNA

Two other families of frogs found in Queensland and the Northern Territory do not occur in Western Australia. The Microhylidae is a group of 19 species of tiny frogs in northeastern Queensland, 1 species in the Northern Territory and a major radiation in New Guinea. The Ranidae is an agile group of pond frogs found on most continents. They are well-represented in New Guinea, but in Australia only one species occurs in Cape York and Arnhem Land (Rana daemeli). Fossil frogs of various ages have been found at numerous sites throughout Australia. In Western Australia fossils of an age of no more than 30,000 years have been found at Skull Cave and Devil’s Lair in the southwest corner. The species Litoria adelaidensis, Crinia georgiana, Limnodynastes dorsalis and Pseudophryne guentheri have been identified; other remains possibly attributable to Litoria cyclorhyncha/moorei, Heleioporus and Neobatrachus have also been recovered but cannot be identified precisely owing to close anatomical similarities between taxa (Tyler 1985).

Hylidae: Litoria coplandi (P. Doughty)

Limnodynastidae: Lymnodynastes depressus (P. Doughty)

Myobatrachidae: Crinia pseudinsignifera (G. Gaikhorst)

Bufonidae: Bufo marinus (J. Francis)

11

Basic frog biology SIZE, FORM AND FEATURES Species of frogs differ substantially in the maximum size that they attain, in the proportions of body and limbs, and in the presence or absence of a number of features. Here we describe and illustrate the nature of these differences. The meaning of some other anatomical and ecological terms is explained in the Glossary (p. 138). Within any species there is variation in adult size. Female frogs are almost always larger than males (a form of sexual dimorphism), and there is also a degree of difference between adults of the same sex. For this reason we quote the range of size for each sex. We refer to size as the length from snout to vent (abbreviated as S-V) measured in millimetres. Another measurement used to quantify the differences between species is the length of the tibia (TL, measured as shown in Figure 1). The TL is used to compute the TL/S-V ratio. If the hindlegs are long, the TL/S-V ratio is high; conversely, a low TL/S-V ratio is an indication of short hindlegs. Extremes are Litoria nasuta which can have a TL/S-V ratio of as much as 0.82, and the stumpy-legged Arenophryne species with ratios as low as 0.14. The eyes of frogs tend to be conspicuous, and the shape of the pupil when constricted is consistent in each species. Generally the pupil forms a vertically or a horizontally orientated ellipse (Figure 2). In genera such as Uperoleia the shape is roughly rhomboidal (Figure 4B). The pupil of Litoria rothii has a horizontal orientation, but there is a V-shaped nick in its lower margin.

Figure 1

External features and methods of measurement of adult frog.

Figure 2

Shape of a frog’s pupil when constricted: (A) vertical slit in Neobatrachus pictus; (B) horizontal ellipse in Litoria caerulea.

a 12

b

BASIC FROG BIOLOGY Figure 3

Variation in the form of the tympanum: (A) annulus sharp in Crinia bilingua; (B) annulus indistinct in Limnodynastes dorsalis; (C) tympanum lacking in Pseudophryne species.

a

b

c

Behind the eye, the external ear (tympanum) may be visible or covered (Figure 3). At its greatest extent the tympanum can be a vast, circular disc almost as large as the eye (see the photograph of Limnodynastes lignarius), but it is generally one-half to three-quarters of the eye diameter. The upper margin of the rim or annulus around the tympanum can be hidden beneath a fold of skin extending from the posterior corner of the eye to a position above the insertion of the forelimb (Figure 3A). This fold is termed the supratympanic fold. Quite often it is difficult to detect the tympanum because the annulus is poorly defined and the skin covering is loose (Figure 3B). In some species, the tympanum is truly absent with no external landmarks. The profile of the head ranges from a low and elongated shape to a short, abbreviated one. The short head is often described as high because development is upwards, so increasing the distance between the external nostril and the margin of the upper lip (compare Figure 3A with Figure 3B). Similarly, in profile the tip of the snout may be projecting (Figure 3A), sloping (Figure 3B), or evenly rounded (Figure 4A). The nostrils (termed nares) most commonly face outwards, and so can be seen as circular apertures in profile (Figure 4A), or else they open upwards rather like those of a crocodile (Figure 4B). Figure 4

Direction of nostrils: (A) facing outwards in Crinia bilingua; (B) facing upwards in Uperoleia crassa.

a

b

13

FROGS OF WESTERN AUSTRALIA Figure 5

Granular skin of ventral surface of a Litoria species.

The dorsal skin may be completely smooth, or it can bear a varying number of circular and slightly raised wart-like structures termed tubercles, when the skin is said to be tubercular. A pair of outward-curving folds forming the shape of a lyre (a musical instrument similar to a small harp) is found on the upper part of the back in the individuals of some species. Such individuals are said to be lyrate. The ventral skin may be smooth or granular (Figure 5). The skin of all frogs contains both microscopic and macroscopic glands. In some species these glands are remarkably developed and form large structures visible externally. Gland development is most conspicuous in Uperoleia species. The glands directly behind the eyes are termed parotoid, those in the groin inguinal, and those at the end of the body coccygeal (Figure 6A). In Litoria splendida, the entire skin covering the roof of the skull forms a gigantic gland termed the rostral gland. On the side of the head in many species there is a further gland (Figure 6B). Although the greatest portion of the gland is situated behind the angle of the jaws, it commences on the upper lip and is termed the supralabial. On the hindlimbs of Limnodynastes dorsalis there are raised, oval glands upon the dorsal surface of the calf, termed the tibial glands. As with many other dermal glands that have been investigated, their secretions are toxic (Crook and Tyler 1981). Cyclorana australis is characterised by an elongated fold of skin extending along the side of the body from a position near the tympanum to the angle of the groin (Figure 7). This fold is therefore at the junction of the dorsal and lateral surfaces, and hence is termed the dorsolateral fold.

Figure 6

(A) Skin glands in a Uperoleia species; (B) gland behind jaw in Limnodynastes dorsalis.

a

14

b

BASIC FROG BIOLOGY Figure 7

Dorsolateral skin fold in Cyclorana australis running forward from the hip.

The frog hand bears four fingers which, for descriptive purposes, are numbered in roman numerals from the inner to the outer (inner cover; Figure 8). On the undersurface of each joint is a circular pad of tissue termed a subarticular tubercle while larger oval or irregularly shaped tubercles on the palm are called palmar tubercles. Almost invariably the sequence of finger length is constant within a species. The frog foot has five toes, and commonly on the sole there are one or two tubercles corresponding to the position of the metatarsal bones beneath them. The innermost is termed the inner metatarsal tubercle (Figure 9) and, if present, the second (near but not beneath the outer metatarsal) is termed the outer metatarsal tubercle. The metatarsal tubercles are highly developed and often sharp-edged in burrowing frogs. The sides of the toes can bear fringes that are narrow and difficult to detect (Figure 9), or are quite conspicuous (Figure 10B). Figure 8

Palm of hand of Limnodynastes dorsalis.

Figure 9

Sole of the foot of Limnodynastes dorsalis.

15

FROGS OF WESTERN AUSTRALIA

The shape of the fingers and toes is highly variable and is commonly modified according to the habits of the species. Ground-dwelling or burrowing frogs tend to have digits like that depicted in Figure 11A, characterised by a simple, elongated, cylindrical form and the absence of lateral fringes. At the other extreme, climbing species have large expanded discs on the tips, and generally lateral fringes as well (Figure 11B). However, the discs may be less markedly developed and scarcely, if at all, broader than the lateral fringes (Figure 11C). An apparent exception to the rule is Limnodynastes dorsalis and Platyplectrum in which, although burrowing species, the female has broad lateral fringes on two or three fingers. However, these fringes are a specialised adaptation enabling the female to produce a foam nest. Between the toes (and less commonly between the fingers as well) there may be a membrane of webbing. This varies in extent from species to species. That shown in Figure 10A would be described as fully-webbed because it reaches the tips of the toes, either directly or via a particularly broadened lateral fringe. Figure 10

Form of the foot: (A) extensive webbing in Litoria caerulea; (B) no webbing but broad lateral fringes in Crinia bilingua.

Figure 11

Form of digits: (A) cylindrical, lacking disc and lateral fringes in Notaden species; (B) flattened with large disc and broad lateral fringes in Litoria caerulea; (C) flattened with small disc and moderate lateral fringes in L. cyclorhyncha. a

b

a 16

b

c

BASIC FROG BIOLOGY Figure 12

Features within the mouth of Limnodynastes dorsalis. The muscles that close the jaws have been cut to permit the jaws to be opened further than would normally be possible and so increase the area visible in the figure.

Inside the mouth there are a number of features of value in identification, but it is only possible to examine them properly in frogs that have been preserved for study. Teeth may be present on or absent from the upper jaw, but are always absent from the lower jaw (Figure 12). They are often extremely difficult to see in tiny frogs even with the aid of a good microscope. They can be detected more readily by feeling them by running the tip of a pin or fine metal rod along the edge of the jaw to detect the tiny serrations. Vomerine teeth may be present or absent on the palate. Generally they are situated upon small mounds projecting downward from the vomerine bones. The vomerine teeth lie between or slightly behind the internal nostril openings (choanae); the tooth rows are most highly developed in the genera Limnodynastes and Platyplectrum. Other apertures visible inside the mouth cavity are the eustachian tubes leading to the inner ears, the slit-like entry to the larynx, the end of the short oesophagus and, in males, short, paired apertures in the floor of the mouth on each side of the tongue leading to the vocal sac. During calling, air passes through the vocal sac slits to inflate the sac located directly beneath. Figure 13

Form of the tongue: (A) broadly cordiform in Limnodynastes dorsalis; (B) oval in Notaden sp.

a

b 17

FROGS OF WESTERN AUSTRALIA

a

b

c

The shape of the tongue falls into one of two categories. The first is that illustrated in Figure 13A, a broad and roughly heart-shaped (cordiform) structure, or else one of a much more oval form (Figure 13B). These differences are much more obvious in observations on living frogs. Then the oval form can be seen to extend for great distances to pick up tiny prey such as ants or termites, whereas the cordiform tongue is far less mobile. As well as possessing vocal sacs, male frogs usually possess nuptial pads upon their first fingers, and sometimes upon the second as well. Some of these structures are unpigmented, glandular structures; others are darkened with keratin (Figure 14B-C). In Heleioporus (Figure 14A) and Limnodynastes lignarius they are replaced by a small cluster of greatly enlarged, sharp, black spines or curved thorns. In several species, the male frog has substantially larger forearm muscles than the female. Examples are Crinia georgiana, Limnodynastes lignarius and Heleioporus (except H. eyrei). Even the position of the vent (cloaca) varies between species (Figure 15), occupying a low position relative to the thighs in Uperoleia (and often being covered by a flap of skin), or being much higher in position and lacking such a flap in Litoria. Figure 15

Direction of opening of cloaca: (A-B) posterior and lateral views of a Uperoleia species in which the cloaca is low relative to position of thighs; (C-D) posterior and lateral views in a Litoria species in which the cloaca is high relative to position of thighs.

18

a

c

b

d

BASIC FROG BIOLOGY

EGGS AND SPAWN The frog egg consists of a spherical ovum surrounded by a mass of clear jelly and enclosed within an outer capsule. Between the ovum and the capsule are two or more additional membranes. The term spawn generally is used to describe a group or clump of eggs. In Western Australian frogs, the diameter of ova ranges from 0.8 mm in Litoria microbelos to at least 5 mm in Myobatrachus gouldii. In general there is not a strong association between the size of the species and egg diameter; larger frogs do not necessarily produce larger eggs. Instead the egg size reflects the lifestyle of the tadpoles or young. A small egg provides a minimal yolk resource and the tadpole hatches quickly and begins to forage for food. In contrast, large egg size is associated with delayed emergence of the tadpole, so that the tadpole is at a more advanced stage when it commences its independent life (as in Pseudophryne species). Indeed, the largest egg size is attained by the species that have no free-swimming tadpole stage, all development occurring within the confines of the outer capsule (the lineage comprised of Arenophryne, Metacrinia and Myobatrachus). The form of the spawn can vary considerably. In the table below we have summarised the diversity in Western Australian species.

Spawn and spawning sites Genus

Spawn

Spawning site

Arenophryne

clumps of large, separate eggs

beneath surface of sandhills

Bufo

laid in long strings

submerged in open water

Crinia

laid singly or in clumps

in open water

Cyclorana

large clump

floating or submerged in open water

Geocrinia

clump

in terrestrial nest or on vegetation above water (G. leai)

Heleioporus

foam nest

burrow in soil

Limnodynastes

foam nest

floating on surface of open water

Litoria

laid singly or in clumps

submerged in open water

Metacrinia

clumps of large, separate eggs

on ground beneath leaf litter

Myobatrachus

clumps of large, separate eggs

beneath surface of sandy or loamy soil

Neobatrachus

string-like, connected chain

submerged in open water

Notaden

large clump

submerged in open water

Pseudophryne

laid singly or in clumps

shallow depressions or water

Platyplectrum

thin foam nest

on surface of open water

Spicospina

laid singly or in small clumps

shallow depressions or channels in peat bogs

Uperoleia

in small clumps

submerged in open water

19

FROGS OF WESTERN AUSTRALIA

TADPOLES AND THEIR MORPHOLOGY Tadpoles are almost as variable in their size and form as adult frogs, and their differences reflect the lifestyles to which they are adapted. Limnodynastes dorsalis represents a tadpole with minimal adaptations to any particular aquatic specialisation, and so it is an ideal subject for a comparative discussion of tadpole structure. As shown in Figure 16, the major division of the tadpole is the separation of the oval head and body from the flattened tail. There is no external division of the head from the body, and internally the entire area is dominated by the coiled intestines, often appearing to be jet black from the nature of their contents visible through the thin body wall. At the front of the head, the feeding apparatus is in the form of a rather mobile funnel lined with several rows of fine, black, comb-like labial teeth above and below the jaws, which resemble two oval, black crescents (Figure 17). Behind the mouthparts is a chamber housing the gills and communicating to the outside by a single aperture (spiracle) on the left side. The anus at the end of the elongated gut is in the midline of the body or to one side of the midline, just where the body joins the tail. The tail is composed of a thin membrane divided into an upper (dorsal) and lower (ventral) fin, and separated by a series of V-shaped muscular blocks which may extend on to the dorsal surface of the body. In general, the species that occupy static water (lentic) tend to have large, deep fins, whereas the inhabitants of flowing water (lotic species) have very low fins, short bodies and extremely muscular tails (see Figure 18). Figure 16

External features and methods of measurement of tadpole of Limnodynastes dorsalis.

Figure 17

Tadpole mouthparts: (A) Limnodynastes dorsalis; (B) L. lignarius.

a 20

b

BASIC FROG BIOLOGY Figure 18

Lateral view of tapoles: (A) Limnodynastes dorsalis; (B) Cyclorana australis; (C) L. lignarius. a

b

c

Litoria moorei tadpole (BB Wells/DEC)

Legislation applying to frogs in Western Australia All native Western Australian frogs are protected under the Wildlife Conservation Act 1950 (WA) (as amended) and the Wildlife Conservation Regulations. The Western Australian Department of Environment and Conservation (DEC) is responsible for the administration of the Act. DEC issues permits for scientific collecting or for zoo displays including import and export permits, and takes special measures to protect the three threatened southwestern species (e.g. Geocrinia alba, G. vitellina and Spicospina flammocaerulea). The Cane Toad (Bufo marinus) is a strictly prohibited species. Recent changes in legislation allow hobbyists to keep frogs as pets. See DEC’s Naturebase website (www.naturebase.net) for more information about what species can be kept and how to go about obtaining a license to keep them. For the enthusiast, the Perth-based Western Australian Herpetological Society meets regularly to discuss herpetology and herpetoculture (www.wahs.org.au). 21

Regional comparisons Examination of the distribution patterns of Western Australian frogs demonstrates that by far the greatest diversity occurs within the Kimberley region, with over 40 species. The frogs of the Kimberley are a clearcut faunal unit, for only six of the species there are shared with the adjacent arid zone, with four of those (Cyclorana australis, C. longipes, Litoria caerulea and L. nasuta) still relying on monsoonal rains in summer for reproduction. Fourteen Kimberley species appear to be restricted to that area (not occurring very far into the Northern Territory, if at all). At a larger scale, the Kimberley shares many species with tropical Northern Territory and Queensland, with several species also occurring in southern New Guinea. The southwest region, like the Kimberley, is an isolated wet corner of Australia with a large endemic fauna. There are over 30 species in the area and, except for a few widely distributed arid zone species (e.g. Neobatrachus sutor), all are endemic to this region. Some taxa have eastern counterparts with no intervening populations in the dry Nullarbor region (Crinia, Geocrinia laevis species-group, Heleioporus, Limnodynastes dorsalis species-group, Litoria aurea species-group, Pseudophryne). The three genera with direct-developing eggs (Arenophryne, Metacrinia, Myobatrachus), the Geocrinia rosea species-group, Spicospina and Litoria adelaidensis are endemic to the southwest region. In the arid zone, many species are widely distributed and extend well into the neighboring eastern states, such as the Cyclorana and Neobatrachus species, and Platyplectrum spenceri. Within Western Australia, many arid-adapated species occur on the edges of the Kimberley and southwest regions, but generally do not penetrate too far into those regions, being replaced by more mesicadapted species. Litoria rubella is an exception, for it is a ubiquitous species in the Kimberley and Pilbara regions. Four species are confined to the Pilbara and Gascoyne regions (Neobatrachus fulvus, Pseudophryne douglasi, Uperoleia glandulosa and U. russelli).

KIMBERLEY

Figure 19

(summer rainfall)

Division of Western Australia into three zones on the basis of rainfall.

ARID ZONE (low irregular rainfall)

SOUTH WEST (winter rainfall)

22

A REGIONAL Approach to identification There are two possible approaches to the identification of the fauna of a geographic unit such as Western Australia. The first is to treat the entire fauna as a single unit, produce a key to distinguish the constituent genera and create a series of separate keys to permit identification of the members of each of the genera. The major disadvantage of this sort of approach is that, whereas at any given locality there may be no more than two or three members of a particular genus to distinguish, identification using a general key can involve comparison with many more species, including some members not found within 2000 km of that locality. The purpose of keys in a field guide such as this is to render identification as simple and accurate as possible when in the bush. Although the strict taxonomic approach of distinguishing each of the members of a genus from each other is useful in other contexts (e.g. when working with museum specimens), this approach can greatly complicate identification in the field. This is because members of a genus are most closely related to one another, thus the characters used to distinguish species can be very subtle. The problem of identification can be reduced, however, if comparisons in a key are restricted to those species occurring in one of the three major Western Australian regions, especially when the genera occur across more than one region (e.g. Crinia, Cyclorana, Litoria and Uperoleia). On a geographic and climatic basis the frog fauna of Western Australia can be divided into three regions (Figure 19): (a) The tropical Kimberley region in the north: extensive rocky country (especially sandstone), woodlands and savannah that receive heavy rainfall during summer and with mild dry winters. (b) The southwest region with a Mediterranean climate: hot dry summers and reliable rainfall during cold winters. (c) The arid zone: a vast landmass situated between these extremes of the state and experiencing low irregular rainfall, usually associated with summer cyclones moving inland from the north. We have adopted the geographic/climatic approach owing to limited overlap between the faunal elements among regions. The arid zone key covers areas away from the southwest and Kimberley regions. If you are working near the boundary of a region and have difficulty identifying a frog, try the key for the adjoining region (the record may also be a range extension near these areas). It remains extremely difficult to distinguish closely-related species that occur together in the same faunal regions, and we draw attention to the use of biological characters in the keys, such as the male advertisement call to distinguish some Crinia species occupying the southwest region, or Uperoleia in the Kimberley region. Difficulty will also be experienced when identifying some Heleioporus, Litoria and Neobatrachus species. In such situations we recommend the use of all data presented here, including known distribution patterns. As a further aid to identification we include checklists of the frog species known to occur within 50 km of each of 12 major cities and towns throughout Western Australia (see pages 136–137 and maps in back cover for locations).

Accounts of fauna Most of the localities cited in the distribution sections of the following accounts are included in the map on back cover. Distributions are defined in terms of the three geographic zones shown in Figure 19. Taxa are presented in alphabetical order with genus first, followed by species. An index of common names is provided at the back of the book. 23

Genus ARENOPHRYNE Tyler, 1976 Diversity and Evolutionary Relationships A genus of myobatrachine frogs endemic to the mid-west coastal region of Western Australia. The sister genus is Myobatrachus, with which it shares a number of osteological features, the habit of head-first burrowing with the hands and a reproductive biology based on direct-developing eggs laid deep in sand.

Morphological Characteristics Body robust. Head blunt with thickened skin on tip of snout; eyes small with horizontal pupil; upper jaw not toothed; covered tympanum marked by fold; tongue long and narrow. Limbs short; fingers short, broad and unwebbed; fourth finger reduced to stump; toes unwebbed.

Reproductive Mode Direct development; i.e. there is no free-swimming tadpole stage as all embryonic stages are passed entirely within the egg and hatchlings are miniatures of the adults. Eggs are laid deep in moist sand.

Etymology areno means ‘sand’ and phryne means ‘toad’.

Further Reading Evolutionary relationships – Maxson and Roberts (1985), Read et al. (2001), Frost et al. (2006); population genetics – Edwards (2007).

Arenophryne rotunda, False Entrance Well (B. Maryan) 25

FROGS OF WESTERN AUSTRALIA

Arenophryne rotunda Tyler, 1976 Northern Sandhill Frog

!! ! ! ! ! ! ! !!

False Entrance Well (B. Maryan)

Description. A small burrowing frog with a squat body; the head is evenly rounded when viewed from above. The limbs are exceptionally short (TL/S-V 0.14–0.22); the fingers are very short, the fourth particularly so, being reduced to a broad stump. The skin surrounding the hind legs is loose, as though several sizes too large. The toes are short and bear numerous prominent tubercles on their undersurfaces. The dorsal colour varies from a uniform off-white or dull cream to grey or light brown, marked with small areas or spots of green, dark brown or brick red. Individuals can change rapidly from a pale to a dark phase. Males 27–31 mm; females 27– 39 mm S-V. Distribution. Southwest region. Shark Bay area, including Edel Land and Dirk Hartog Island. Habitat. Found among coastal sand dunes. This species uses its hands to burrow and during the rainy winter months emerges nightly from a cell 10 cm below the surface. It can be detected by the tracks that it leaves upon the surface of the sand. Advertisement Call. A short ‘squelch’ emitted underground or on the sand surface.

26

Breeding Biology. Males call from July to November. Pairing takes place during this time, and couples remain together until late March to April when the eggs are laid. Clutches of 4–11 large (5–6 mm) creamy white eggs are laid at a depth of 80 cm in moist sand. There is no freeliving tadpole stage: the embryonic stages are passed entirely within the egg capsule. Hatching takes place about 2 months later and hatchlings are about 10 mm. Etymology. rotunda means ‘round’ or ‘spherical’ in reference to the shape of this species. Remarks. This species was listed as ‘Threatened’ for some years after its discovery and description in 1976 as so few encounters had been reported. However, it was subsequently found to be extremely abundant under favourable conditions and employing pit-trapping capture techniques (Roberts 1985). Further Reading. Call analysis – Roberts (1984); ecology – Tyler et al. (1980), Roberts (1990); larval development – Roberts (1984), Anstis et al. (2007); skeleton – Davies (1984); physiology – Cartledge et al. (2006a); population biology – Roberts (1985).

Genus ARENOPHRYNE

Arenophryne xiphorhyncha Doughty and Edwards, 2008 Southern Sandhill Frog

! ! ! ! !! !!

Cooloomia Outstation (B. Maryan)

Description. A small to medium-sized frog with a flattened oval-shaped body and short limbs (TL/S-V 0.14–0.24). The skin is loose with slightly raised bumps and folds on the back, tending to form longitudinal ridges. The eyes and snout are smaller than in A. rotunda, but the hands are larger. The dorsum ranges from a creamy yellow to dark brown flecked with red with a thin pale vertebral stripe. Males 27–32 mm; females 26–36 mm S-V. Distribution. Southwest region. Central west coast from Kalbarri National Park north and inland to Cooloomia station. Habitat. Occurs on much coarser sands than A. rotunda, possibly explaining its smaller, more streamlined head and larger hands which would assist in burrowing forwards through the sand.

Advertisement Call and Breeding Biology. Unknown but probably similar to A. rotunda. Etymology. xiphos means ‘sword’, and rhynchos means ‘nose’ or ‘snout’; thus, ‘sword-snouted’ for this spieces in contrast to the more broadlyrounded snout of A. rotunda. Remarks. This new species was discovered as a result of genetic tests run by D. Edwards, and subsequently confirmed by examining museum specimens from Kalbarri to Shark Bay. The description of a second species of Arenophryne means each species is confined to a small natural range. Further Reading. Population genetics – Edwards (2007).

27

Genus CRINIA Tschudi, 1838 Diversity and Evolutionary Relationships A diverse genus of myobatrachine frogs. They are relatively distantly related to other myobatrachines, but are probably closest to Geocrinia, Assa and Paracrinia. There are a total of 15 species distributed across most of Australia except the arid centre and Pilbara region. They also occur in southern New Guinea. Five species occur in the southwest and three in the Kimberley.

Morphological Characteristics Small body size, with most species 2–3 cm long. Head and body flattened; upper jaw toothed; tympanum present but covered; tongue long and oval. Limbs short; fingers and toes long and unwebbed; toes slightly fringed. Many species display polymorphism in back pattern with three morphs: smooth (no longitudinal folds or markings), lyrate (a pair of folds near the shoulders) and ridged (continuous longitudinal folds and markings).

Reproductive Mode All Crinia species lay eggs in shallow water. Eggs are pigmented and usually laid singly, typically in small clusters (except for C. riparia which lays large clutches under stones in streams, and C. nimbus with a terrestrial nest). Larval life is extended (> 3 months) in most species, with tadpoles attaining large size relative to the adult frog. Crinia georgiana is unusual for its more rapid development (30–45 days to metamorphosis) and the tiny size of the metamorphs. Crinia bilingua and C. fimbriata are also capable of rapid development as these species breed in summer.

Etymology Uncertain, but possibly related to ‘lillypad’ or perhaps just an arbitrary name coined for its euphonius sound.

Remarks Owing to the highly derived breeding biology of C. georgiana (the type species of Crinia), the remaining smaller-bodied Crinia were once referred to the genus Ranidella Girard, 1853. Recent genetic work indicates that Crinia forms a natural group, with C. georgiana closely related to small-bodied southwest species (Read et al. 2001; Frost et al. 2006).

Further Reading Evolutionary relationships – Barendse (1984), Read et al. (2001), Frost et al. (2006).

Crinia glauerti, Walpole (G. Harold) 29

FROGS OF WESTERN AUSTRALIA ! ! !! !!! ! ! ! ! ! !! ! ! !! ! ! ! !! !!!! ! !! ! !! !! !!!! ! ! !! ! ! ! !! ! ! !

Crinia bilingua (Martin, Tyler and Davies, 1980) Bilingual Froglet

Kununurra (P. Doughty)

Description. A small, slender-bodied pale brown or grey frog with or without an elaborate pattern of bars and stripes across the body and limbs. The limbs are of moderate length (TL/S-V 0.37–0.50) and slender. The fingers and toes are very long, slender and unwebbed, but the toes bear well-developed lateral fringes (Figure 10). Males 16–24 mm; females 17–29 mm S-V. Distribution. Kimberley region. West to Yampi Peninsula and south to Mount Barnett, Tableland and the Black Elvire River. Also occurs in the Northern Territory. Habitat. Streamsides and areas where water persists during the dry season. Advertisement Call. A high-pitched rattle or clicking sound with a musical quality. The call

30

has two parts, hence the scientific name. Breeding Biology. This species calls from the base of grass tussocks at the edge of swamps and temporarily flooded areas. There is evidence that this species is an opportunistic breeder, likely to breed whenever suitable conditions occur. Tadpoles grow up to 26 mm in total length and larval life may be as short as 2 weeks owing to high temperatures experienced in summer for this species compared to the southwestern species that breed in winter. Etymology. bilingua refers to the two-part (‘twovoiced’) advertisement call. Futher Reading. Call analysis – Martin et al. (1980).

Genus CRINIA

Crinia deserticola (Liem and Ingram, 1977) Chirping Froglet

! !

Thargomindah, QLD (H. Cogger)

Description. A small, slender-bodied pale grey species with limbs moderately stout and long (TL/S-V 0.37–0.48). The fingers and toes are long, slender and unwebbed. There are two main dark blotches on the back. Males 13–18 mm S-V. Distribution. In Western Australia only known from near Lake Argyle. It occurs in arid regions in the Northern Territory and Queensland, extending as far as Townsville to the northeast. Habitat. Associated with blacksoil plains and mulga.

Advertisement Call. A high-pitched chirping, like a sparrow. Breeding Biology. Breeds along creeks and flooded areas after summer rains. Etymology. deserticola means ‘desert-dweller’. Remarks. Distinguished from C. bilingua in having the dorsal pattern disrupted into smaller blotches. Further Reading. Call analysis – Liem and Ingram (1977), Tyler et al. (1981c).

31

FROGS OF WESTERN AUSTRALIA

Crinia fimbriata Doughty, Anstis and Price, 2009 Kimberley Froglet

! !

Little Merten’s Falls (L. Price)

Description. A small-bodied species with relatively long legs (TL/S-V 0.48–0.54). Males have wide flanges on the fingers; belly smooth. Background colour is a bluish or reddish greybrown with a network of fine bluish-white tubercles scattered on the back. Males 16–17 mm S-V. Distribution. Kimberley region. Only known

32

from the Mitchell Plateau and Prince Regent River region in the northwest Kimberley. Habitat. Sandstone platforms in highly dissected country. Advertisement Call. Unknown. Breeding Biology. Males have been found in shallow rock pools on sandstone outcrops in summer. The tadpoles are small (to 20 mm) with a wide body and truncate snout. The jaws are highly developed and may enable tadpoles to scrape algae from the bottom of rock pools where they have been found. Etymology. fimbriata means ‘fringed’, in reference to the flanges on male fingers. Remarks. Known from only a few individuals, the call and the function of the flanges on the fingers of males are unknown.

Genus CRINIA

Crinia georgiana Tschudi, 1838 Quacking Frog

! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! !! ! !! ! ! !! ! !! !! ! ! ! !! ! !! ! ! !!! ! ! ! ! ! !!! ! !!! ! ! ! ! !! ! ! ! ! ! !!!! ! !!!! ! ! ! ! ! !! !! !! !! ! ! ! ! ! ! !!!! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! !! !! ! ! ! !!! ! !!

!!! ! !!! ! !! !

Albany (Greg Harold) (L) ♂ Northcliffe (B. Maryan) (R) ♁ Walpole (B. Maryan)

Description. The largest species of Crinia. A squat and comparatively flattened species with a rather large head, short limbs (TL/S-V 0.36– 0.41) and unwebbed fingers and toes. Males have massive arms. The dorsal skin ranges from smooth (most females) to tubercular; the abdomen is granular. Colouration of markings upon upper parts of body is highly variable through pale grey, brown to red, often with an elaborate pattern, to almost black. All individuals have a bright red patch in the groin and red or golden patches on the upper eyelids. The ventral surface of females is stark white, but variable in males. Males 19–44 mm; females 28–41 mm S-V. Distribution. Southwest region. Southwest from Gingin in the north inland to Dumbleyung and east to Cape Le Grande. Also on Mondrain Island.

Habitat. Occurs in a variety of habitats, especially woodlands and near granite outcrops. Breeds in low-lying areas or slow-flowing seepages with only very shallow water (1–5 cm deep). Advertisement Call. A short and loud call, resembling the quack of a duck. Breeding Biology. This species has an atypical and interesting reproductive life which has been the subject of much recent work at the laboratory of J. D. Roberts (University of Western Australia). Breeding occurs from July to October on cold nights. In a large proportion of matings, several males will simultaneously embrace the female (and often each other) in order to fertilize her eggs. The large arms of the males help them to wrestle for a good position on top of, and underneath, females to achieve some fertilization

33

FROGS OF WESTERN AUSTRALIA

success. Females lay from 90 to nearly 300 eggs which she releases in about 10 minutes. The eggs are large and separate and are often laid in shallow water from seepages and roadside ditches. Tadpoles show lotic adaptations and complete metamorphosis at a tiny size in 30–45 days. Tadpoles can accelerate metamorphosis in response to pond drying or food shortages. Etymology. After King George Sound off Albany, the type locality for the species.

Further Reading. Calling behaviour – Ayre et al. (1984), Gerhardt et al. (2000), Smith and Roberts (2003a,b), Smith et al. (2003a); mating behaviour – Roberts et al. (1999), Smith and Roberts (2003a,b), Byrne and Roberts (1999, 2000, 2004), Byrne (2002, 2004), Dziminski et al. (2008); developmental biology – Main (1965a), Seymour and Roberts (1995), Seymour et al. (2001), Doughty (2002), Smith and Roberts (2003c), Doughty and Roberts (2003), Dziminski and Roberts (2006); population genetics – Edwards et al. (2007).

Figure 20

Characteristic ventral patterns: A) male Crinia insignifera; B) female C. insignifera; C) male C. glauerti; D) female C. glauerti.

a

B

C

D

34

Genus CRINIA

Crinia glauerti Loveridge, 1933 Rattling or Clicking Froglet

! !! !! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! !! ! ! ! ! !! ! !!!! !! ! ! !! ! ! !!! ! ! ! ! ! ! !! !!! !!!! !! ! ! ! ! !!! !! ! ! !! ! !!

Walpole (G. Harold) Denmark (B. Maryan)

Description. A highly variable, small species with moderately long legs (TL/S-V 0.39–0.47). The dorsal skin is smooth or rough and has a range of patterns of lines and patches of grey, brown, black and white as in other Crinia. The female has large conspicuous black markings upon the undersurface (Figure 20D), and these sometimes occur in males. Males are uniformly black on the throat, often extending on to the upper abdomen (Figure 20C). Males 10–22 mm; females 14–24 mm S-V. Distribution. Southwest region. North to the Moore River; south and east to the Pallinup River; inland to Chittering, Dwellingup, Green-

bushes and the Stirling Ranges. Advertisement Call. An irregular rattling that trails off. Main (1965a) compares it to a ‘long hollow rattle like a pea in a can’. Habitat. Areas of permanent or seasonal moisture such as the edge of marshes and near rocky outcrops. Breeding Biology. Other than in summer, this species will breed following any rain. The eggs sink to the bottom of swamps and static areas at the edge of streams. Tadpoles are very heavily pigmented and take more than 3 months to metamorphose into frogs. Main (1965a) reports that members of this species can be sexually mature at only 12–13 mm length. Etymology. Named for Ludwig Glauert (1879–1963), director of the WA Museum who described several species of Australian frogs and reptiles. Further Reading. Call analysis – Littlejohn (1959), Littlejohn and Wright (1997); diet and biology – Main (1957a, 1965a, 1968).

35

FROGS OF WESTERN AUSTRALIA

Crinia insignifera Moore, 1954 Squelching Froglet

! ! ! ! ! ! ! ! ! ! ! ! ! !! !! ! ! ! ! ! ! ! ! ! ! !! ! ! ! !!! !

Capel (G. Harold)

Description. A small, long-limbed frog (TL/S-V 0.40–0.53) with unwebbed fingers and toes. The skin of the back may be smooth, lyrate or have longitudinal ridges. The colour ranges from brown to grey without markings or with various patterns of longitudinal stripes or patches. A triangular dark patch is commonly found upon the head directly between the eyes, and the legs often bear dark transverse bars. The abdomen in both sexes is marked with irregular black or grey blotches. In breeding males the edge of the lower jaw is darkly pigmented (Figure 20A). In females the throat is unpigmented (Figure 20B). Males 14–23 mm; females 16–30 mm S-V. Distribution. Southwest region. The coastal plain between Gingin and Bussleton. A remnant population persists on Rottnest Island. Habitat. Permanent and seasonal wetlands of the Swan Coastal Plain. Advertisement Call. A high-pitched squelch. Described by Main (1965a) as sounding ‘like drawing a wet finger over a balloon’. Breeding Biology. A winter breeder. Eggs are laid singly in pools and fall to the bottom.

36

The tadpoles take up to 5 months to complete development. Etymology. A reference to C. signifera (‘signbearing’) from southeastern Australia owing to the small morphological differences between the species; thus, ‘inside signifera’. Remarks. Morphologically, this species is almost indistinguishable from the closely-related C. pseudinsignifera; however, the two species can be easily separated on calls. They hybridise near the base of the Darling Scarp and the hybrid zone was examined by several investigators from the late 1950s to the early 70s. The Rottnest Island population was believed to be extinct owing to roadworks during the 1940s and 50s that destroyed most of the freshwater lakes on the island. It is still present but rare (C. Wheatley, pers. comm.). Further Reading. Call analysis – Littlejohn (1959); reproduction – Main (1957a, 1968); inheritance of back patterns – Main (1961, 1965b, 1965c), Bull (1975); hybrid zone – Blackwell and Bull (1978), Bull (1978, 1979); Rottnest Island population – Main (1961, 1965b).

Genus CRINIA

Crinia pseudinsignifera Main, 1957 Bleating Froglet

! !! ! ! !! ! ! ! ! ! ! !!!! ! !! ! ! !!!! ! ! ! !! ! !! ! ! !! ! ! ! !! ! ! !! !! ! !! ! ! ! ! ! !! ! ! !! !! ! ! !! ! !! ! ! ! !! ! ! ! ! ! ! ! ! !! !! !! ! ! !! !! ! !!!! !!! ! ! !!!!!!! ! ! ! ! ! !!!! ! !! ! ! ! ! ! !!! !! ! ! ! ! ! !! ! ! ! ! ! !!! !!! !! ! ! !! ! !!!! ! !! !

! ! !

Ponier Rock (B. Maryan)

Description. A small long-limbed frog (TL/ S-V 0.43–0.51) with unwebbed fingers and toes. The back can be smooth, bumpy or have longitudinal ridges. In its external features it is not readily distinguishable from C. insignifera, but can be identified by the advertisement call and distribution. Males 14–24 mm; females 18–29 mm S-V. Distribution. Southwest region. Generally associated with granites on the Pre-cambrian Shield south of a line joining the lower Murchison River (near Kalbarri), Bonnie Rock, Bank Rock and Nanambinia. Absent from the Swan Coastal Plain where it is replaced by C. insignifera. Habitat. Seasonal wetlands and waterways and associated with granite outcrops in the wheatbelt.

Advertisement Call. A high-pitched wavering call of 3 or 4 pulses repeated frequently, sounding somewhat like the bleating of a sheep or goat. Breeding Biology. Breeds in winter, with reports of summer breeding at the arid edge of their range (Lake Cronin). Eggs are laid singly and have been found beneath cover in waterlogged soil adjacent to pools (Main 1965a). Larval life ranges from 80 to 130 days. Etymology. Meaning ‘false C. insignifera’ owing to the similarity to that species, which had only been described 3 years previously by J. A. Moore. Further Reading. Call analysis – Littlejohn (1959); tadpole description – Main (1957a); hybrid zone – see C. insignifera.

37

FROGS OF WESTERN AUSTRALIA

Crinia subinsignifera Littlejohn, 1957 South Coast Froglet

! ! !!! ! !! ! ! !!

!

!

Northcliffe (B. Maryan)

Description. In its external features similar to C. insignifera and C. pseudinsignifera from which it can be distinguished by the greater degree of lateral fringing of the toes. The limbs are moderately long (TL/S-V 0.40–0.50). Males 14–20 mm; females 20–24 mm S-V. Distribution. Southwest region. From Manjimup, Frankland, Stirling Ranges and Pallinup River in the north, south to Pemberton, Lake Muir and vicinity of Albany; that is, a narrow band across the northern edge of the karri belt. Also on the south coast at Bremer Bay, Fitzgerald National Park, Cape Le Grand and Cape Arid. Habitat. Permanent or seasonally inundated Melaleuca swamps.

Advertisement Call. A drawn out low-pitched squelch that fades away. Breeding Biology. Males call from around submerged sedges in relatively deep water. Also observed breeding in shallow roadside ditches. Sympatric with C. pseudinsignifera where both species will breed in the same ponds. Etymology. Meaning ‘beneath C. insignifera’ referring to the range of this species occurring to the south of C. insignifera. Further Reading. Evolutionary relationships – Read et al. (2001); call analysis – Littlejohn (1959).

Cyclorana australis, Kununurra (P. Doughty) 38

Genus CYCLORANA Steindachner, 1867 Diversity and Evolutionary Relationships A moderately diverse genus of burrowing hylid frogs, with a total of 12 species distributed across the central arid regions and tropical north of Australia. Seven species occur in Western Australia. This genus was at one time included within the Leptodactylidae (i.e. Limnodynastidae and Myobatrachidae), but belongs within the Hylidae on morphological and biochemical grounds (Tyler 1972, 1979; Maxson et al. 1982, 1985; Frost et al. 2006). The closest relatives of the genus Cyclorana are species of the Litoria aurea species-group, such as L. cyclorhyncha (Tyler et al. 1978; Hutchinson and Maxson 1986; Burton 1996). Frost et al. (2006) only recognize Cyclorana as a subgenus within Litoria, but we adopt the more traditional arrangement here pending further taxonomic revision of the Hylidae.

Morphological Characteristics Body robust; head high or flattened; eyes prominent; pupil horizontal; mouth wide; upper jaw toothed; tongue broad; tympanum visible or covered. Limbs short; fingers unwebbed; first finger opposable and longer than second; toes long, partly or completely webbed; shovel-shaped inner metatarsal tubercle.

Reproductive Mode Aquatic reproduction. Eggs pigmented; laid in large clumps in water. Development can be extremely rapid, reflecting the temporary availability of water throughout much of the arid and tropical regions.

Etymology Cyclo means ‘round’ and rana means frog; hence, ‘round frog’ in reference to the shape of the first species described (C. australis).

Remarks Some arid-adapted species of Cyclorana can form a protective skin ‘cocoon’ as a means of reducing water loss during long periods of aestivation (Lee and Mercer 1967; Withers 1995a, 1998).

Further Reading General – Tyler and Davies (1993).

FROGS OF WESTERN AUSTRALIA

Cyclorana australis (Gray, 1842) Giant Frog !

! !! ! ! ! !

!! ! ! ! !! ! ! ! ! ! ! ! ! !! !! ! ! ! ! !! ! ! !! ! ! ! !!! ! ! ! ! ! ! ! ! ! !! !! ! ! ! ! ! !! ! ! !!! !!! !! !! !! !!!! !!!! !! ! ! ! ! ! ! ! !!! ! !!

Kununurra (B. Maryan) Sandfire Roadhouse (B. Maryan)

(Godfrey Tank) in the east. Also throughout tropical and subtropical Northern Territory and the Gulf Country of Queensland; replaced by the sister species C. novaehollandiae to the east. Habitat. Active at night in open, poorly vegetated areas. Commonly sighted on roads. Burrows about 10 cm below the ground surface during the day, but also known to bask in the sun near water reaching body temperatures of up to 34°C (Tyler 1994). Advertisement Call. A loud ‘whoark’ repeated Description. This is one of the largest native frequently. ground-dwelling frogs in Australia. It is a robust and Breeding Biology. Gravid females carry about 1000 muscular frog with moderately long limbs (TL/S-V eggs. They spawn at the commencement of the wet 0.38–0.50). The head is prominent and triangular, season (October–December) and are one of the first and the eyes large, dark and rather protuberant. frogs to use the newly filled ponds. Tadpoles attain a The dorsal skin is rough, bearing distinct small maximum length of 60 mm. The mouth is pointed tubercles. A diagnostic character is a pair of raised, downwards with 2 upper and 3 lower tooth rows, dorsolateral, glandular skin folds on the back, and the inner of each being narrowly divided medially. often a further pair on the sides (Figure 7). Fingers Tadpoles form large conspicuous schools. are elongate and unwebbed; the feet have long toes, minimal webbing and a pronounced inner metatarsal Etymology. australis means ‘southern’ but in this tubercle. They are commonly pale brown or grey context means ‘from Australia’. above, but occasionally extensively marked with Remarks. Cyclorana australis commonly feeds on green (especially juveniles). A poorly defined dark smaller frogs. In the Northern Territory, C. australis stripe extends along the side of the head from the is the species most commonly mistaken to be a Cane tip of the snout to the eye, and from behind the eye Toad and killed. Cane Toad tadpoles also form large to the angle of the jaw. Males 71–85 mm; females schools, but are much smaller (1–2 cm versus 6 cm) and are black instead of cream. 71–102 mm S-V. Distribution. Kimberley region and extending to the arid zone in the Pilbara (south and inland Further Reading. General – Tyler and Martin (1975); to Woodstock) and the Canning Stock Route cocoons – Withers and Thompson (2000). 40

Genus CYCLORANA

Cyclorana cryptotis Tyler and Martin, 1977 Hidden-ear Frog

! !!! ! ! ! !!!! !! ! ! ! !

Kununurra (P. Doughty)

Description. A moderate-sized and rather globular frog with a relatively small and compressed head that is gently rounded, and not triangular or as prominent as in other Cyclorana species. The tympanum is completely hidden beneath the skin. Limbs are short to moderately long (TL/S-V 0.34–0.40). The skin is only slightly roughened. The fingers are cylindrical and unwebbed and the foot bears a prominent inner metatarsal tubercle. Colour ranges from pale grey in juveniles to brown, lightly mottled with brown in adults. There are 5 wide longitudinal stripes that can be a dull or strikingly bright orange. Some individuals bear a pale, narrow mid-vertebral line. Males 34–46 mm; females 36–48 mm S-V. Distribution. Kimberley region. Southern areas: west to Dampier and Yampi peninsulas; in the east from Wyndham to Lake Argyle. Extends east to the Northern Territory and Queensland.

Habitat. Open grassland. Advertisement Call. A loud modulated ‘wwooaawwrr’ repeated frequently. Breeding Biology. An ‘explosive’ wet season breeder – large choruses may form after heavy rains, with little or no calling or breeding at other times. Males call from the base of tall grasses at the edge of temporary ponds or ditches and often call while floating in shallow water. Spawn is deposited as large irregular masses in murky, colloidal water in shallow pools on clay soils where temperatures above 40°C are common. Tadpoles attain a maximum length of 46 mm. The mouth is subterminal with 2 upper and 3 lower tooth rows. Larval life is about 1 month. Etymology. cryptotis means ‘hidden ear’. Further Reading. General – Tyler et al. (1982); distribution – McDonald (1998).

41

FROGS OF WESTERN AUSTRALIA

Cyclorana cultripes Parker, 1940 Knife-footed Frog

! ! !! !

Kununurra (B. Maryan)

Description. A small to moderate-sized species with cylindrical, unwebbed fingers, half-webbed toes, limbs short to moderately long (TL/S-V 0.33–0.40). Most individuals are pale grey with irregular darker markings of olive green. Occasionally there is a narrow, pale straw-yellow, mid-vertebral stripe, but most individuals have a slightly broader mid-ventral zone devoid of darker markings; there is usually a pale bar behind the eyes. The posterior surface of the thighs is hazel, on which there are small, pale patches of lighter pigment. Males 38–46 mm; females 38–43 mm S-V. Distribution. Kimberley region. Confined to the eastern edge. Widely distributed throughout the southern half of the Northern Territory, southwest Queensland and northern South Australia. Habitat. Floodplains adjacent to creeks and other low-lying areas exposed to flooding in the wet season.

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Advertisement Call. The male call is a very short, harsh note. Breeding Biology. Males call from open positions within 1m of small pools. The tadpole has a long tapering tail tip, a dextral anus, and 2 upper and 3 lower rows of labial teeth. The oral disc is surrounded by papillae except at the upper margin. Etymology. cultripes means ‘knife-footed’. Remarks. Cyclorana cultripes and C. vagitus are extremely similar morphologically, and can only be distinguished on size, call and geography. Main’s (1965a) account of ‘C. cultripes’ in fact refers to C. maini. Further Reading. Call description – Tyler and Martin (1977); tadpole description – Watson and Martin (1973); cocoons – the report of Withers and Thompson (2000) is actually based on C. longipes.

Genus CYCLORANA !! ! ! ! ! ! ! ! !!!

Cyclorana longipes Tyler and Martin, 1977 Long-footed Frog

! ! ! !! !! ! ! ! ! ! ! ! ! ! ! !! ! ! ! ! ! !

! !!!! ! !! ! ! !

!! ! ! ! !

Mornington Station (P. Doughty)

Description. A medium-sized species with a rounded snout, protruding eyes and smooth skin. One of the most strikingly marked of all Cyclorana, it bears a contrasting pattern of clearly-defined, very dark brown or olive markings upon a pale grey background. The side of the head bears a broad dark stripe from the nose through the eye and there is a dark, irregular margin to the upper jaw. The limbs are short to moderately long (TL/S-V 0.36–0.45). The fingers are slender, unwebbed and lack lateral fringes, and the toes are long and webbed mainly at the base. Males 38–51 mm; females 36–55 mm S-V. Distribution. Kimberley region. Extending to the arid zone to the southwest to Anketell Ridge and to Well 46, Canning Stock Route, to the southeast. Also occurs throughout the northern portion of the Northern Territory. Habitat. This ubiquitous species occupies a wide variety of habitats in northern Australia, indicating that it can tolerate a wide variety of soil types.

Advertisement Call. A long, loud, high-pitched rising then falling note repeated frequently. Males in choruses call in unison. Breeding Biology. Breeds at the commencement of the wet season; it is one of the first northern species to breed after rain. Spawn clumps ranging from 50–2000 eggs are laid in shallow water and sink to the bottom. Tadpoles resemble C. australis tadpoles until the last period of development, and complete metamorphosis in approximately 1 month. Etymology. longipes means ‘long-footed’ in reference to the slightly longer feet of this species compared to other Cyclorana. Remarks. Most closely related to C. maculosa of the Northern Territory and C. brevipes of Queensland. Further Reading. Call analysis – Tyler and Martin (1977); cocoon formation (as C. cultripes) – Withers and Thompson (2000).

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FROGS OF WESTERN AUSTRALIA

Cyclorana maini Tyler and Martin, 1977 Sheep Frog or Main’s Frog ! !! ! ! !! ! !! !! ! ! !! !! ! ! ! ! ! !!! ! !! !! ! ! ! ! ! ! ! ! ! !!!! !! ! ! ! !!! ! ! ! !! ! ! ! ! ! !! ! ! !! ! ! ! ! ! ! !!! ! ! ! ! ! ! ! ! ! ! !! !!! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! !! ! ! ! ! ! ! ! !

!

!

!

Leinster (G. Harold) Newman (B. Maryan)

Description. This abundant frog is a moderatesized, rotund species. The fingers are slender and unwebbed and the toes approximately half-webbed. The limbs are short to moderately long (TL/S-V 0.33–0.41). The back is generally dull grey or pale brown and bears small and usually longitudinally-arranged darker brown or olive markings, or else darker markings of indeterminate shape. Some individuals can change their background colouration from green to brown within a few hours. All exhibit a dark stripe from the nostril to the tympanum. Males 35–46 mm; females 39–48 mm S-V. Distribution. Arid zone. Pilbara and Gascoyne regions, south and inland to Mount Magnet,

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Leonora and Lake Yeo; north and inland to Anketell Ridge, Durba Hills and the Walter James Range. Also occurs in the southern portion of the Northern Territory and northern South Australia. Habitat. Most commonly found on floodplains and in areas near temporary stream beds. Advertisement Call. Similar to the bleating of sheep or the noise of an ambulance siren. Breeding Biology. Main (1965a) states that the eggs and early stage tadpoles resemble those of C. platycephala, but that the later stages are pink with an opalescent sheen. Length at metamorphosis is reported to be 19 mm. Etymology. Named after A. R. (‘Bert’) Main from the University of Western Australia, a pioneer of studies of Western Australian frogs. Further Reading. Abundance – Morton et al. (1993); call analysis – Tyler and Martin (1977); diet and reproductive biology (as C. cultripes) – Main and Calaby (1957), Main (1965a), Main and Storr (1966), Calaby (1960); tadpoles – Main (1965a); physiology and cocoon formation – Withers (1993, 1995a, 1998).

Genus CYCLORANA

Cyclorana platycephala (Günther, 1873) Water-holding Frog ! !

!

! ! ! !

! ! ! !

! !

! !

!! ! ! !

!! ! ! ! !

!! ! !

!

! ! ! ! ! ! !!

!

! ! ! !! ! ! !!

Meedo Station (B. Maryan)

Description. A large-bodied species with a flattened appearance and wide mouth. The eyes are small and placed high upon the head. The limbs are relatively short (TL/S-V 0.35–0.42) and the toes are unique in Cyclorana in being fully webbed. The colouration changes with reproductive activity. During most seasons, they are pale grey to brown with marbling of a slightly darker shade, becoming extensively marked with green after periods of activity – usually in the periods preceding spawning. Males 40–47 mm; females 49–68 mm S-V. Distribution. Arid zone. In the north from the North West Cape, southern Pilbara and east to Lake Disappointment, and south to Morawa and Laverton. Also in arid zones of the mainland states (except Victoria). Habitat. Widely dispersed over a variety of environments, including several types of impervious soil such as clay. In dry conditions it burrows to a depth of 1m. A predator of other frogs and tadpoles and one of the last species to burrow when ponds dry.

Advertisement Call. A long drawn-out ‘maw-ww-w-w’. Breeding Biology. Breeds after cyclonic rain. Spawn is laid in large masses of up to 1700 eggs in temporary pools. Tadpoles are large (to 8 cm total length) and development takes about 30–50 days. At metamorphosis the young have a body length of up to 35 mm. Etymology. After the Greek roots of platy meaning ‘flat’ and cephalos meaning ‘head’, hence ‘flat-headed’. Remarks. This is one of the best known of the ‘water-holding’ burrowing frogs of Australia. Further Reading. Distribution – Tyler (1990); call description – Main (1965a); reproduction – Anstis (2002); physiology and cocoon formation – Main and Bentley (1964), Van Beurden (1979, 1982), Withers (1993, 1995a, 1998), Cartledge et al. (2008); field observations – Spencer (1896); distribution – Tyler (1990).

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FROGS OF WESTERN AUSTRALIA

Cyclorana vagitus Tyler, Davies and Martin, 1981 Wailing Frog

! !!!! ! ! ! !! ! ! ! ! ! !

Kununurra (G. Harold)

Description. Similar in appearance to C. cultripes, but larger-bodied. A moderate-sized, robust-bodied frog with a tubercular skin and diffuse grey and dark green mottling on the back. The limbs are short to moderately long (TL/S-V 0.36–0.44). The toes are approximately half-webbed. The male has 1 or 2 nuptial pads on the first finger. Males 42–48 mm; females 44–48 mm S-V. Distribution. Kimberley region. Occurs along the Fitzroy and Lennard Rivers in the west, and the lower Ord valley in the east. Habitat. Occurs on open grassy plains.

Advertisement Call. Resembles the crying of an infant – ‘waaa…waaa’ – repeated slowly and frequently. Breeding Biology. Breeding has been observed in January and February following heavy cyclonic rains. They breed in shallow static water with a depth of 15–30 cm. Etymology. vagitus refers to ‘wailing’ or the crying of an infant in reference to the male’s call. Remarks. This species is closely related to C. cultripes, but differs in call and attaining a larger size.

Geocrinia leai, male on eggs (© Jiri Lochman, Lochman Transparencies) 46

Genus GEOCRINIA Blake, 1973 Diversity and Evolutionary Relationships A small but disparate genus of myobatrachine frogs, with 7 species distributed across southern Australia. Five species endemic to the southwest of Western Australia. Among the southwestern species, G. leai is more closely related to the 2 species from southeastern Australia, with Assa appearing to be the sister taxon.

Morphological Characteristics Head and body broad, rather flattened; upper jaw toothed; tympanum present. Limbs short; fingers and toes unwebbed; toes with or without terminal discs. Ventral skin smooth (all Western Australian species) or granular.

Reproductive Mode Two very distinct modes. In G. leai, eggs laid in clumps attached to aquatic or riparian vegetation above the waterline. Upon hatching, tadpoles drop into water to commence a long period of free-swimming development. In the remaining species, eggs are laid in terrestrial nests (e.g. small depressions or shallow tunnels). Tadpoles hatch from egg capsules but remain in nest until metamorphosis.

Etymology Geo means ‘earth’ (in reference to the site of egg deposition), hence ‘earth Crinia’.

Remarks Two species (G. alba and G. vitellina) are listed as Threatened Fauna, based on their restricted ranges, lack of dispersal and associated degree of genetic subdivision and continuing loss of populations owing to clearing and fragmentation.

Further Reading Evolutionary relationships – Roberts and Watson (1993), Wardell-Johnson and Roberts (1993, 1996), Read et al. (2001), Frost et al. (2006).

FROGS OF WESTERN AUSTRALIA

Geocrinia alba Wardell-Johnson and Roberts, 1989 White-bellied Frog

!!

Forrest Grove (B. Maryan) Witchcliffe (R. Browne-Cooper)

Description. A small frog with limbs moderately developed and short (TL/S-V 0.36–0.40). Very similar to G. vitellina but with a white ventral surface, with or without a faint yellowish wash. Back light brown or grey with darker brown spots, usually centred on pores and almost forming flattened warts, the most prominent of which are aligned, sometimes coalescing to form a stripe from the eye to the cloaca and a short medial row on the rump. Larger brown spots also form a mid-lateral line from eye along flank and a transocular bar. In some individuals there is a pink suffusion on the rump and shoulders. Males 20–24 mm; female 17 mm S-V. Distribution. Southwest region. Known only from about 130 km2 in the Karridale-Witchcliffe area.

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Habitat. Dense vegetation on clay. Advertisement Call. An irregularly repeated discrete series of 11–18 pulses given almost too rapidly to be resolved by ear. Breeding Biology. Males call from small depressions in clay under dense vegetation. Egg masses often associated with calling males. Etymology. alba means ‘white’ in reference to the pale belly. Remarks. Compared to G. vitellina, the dorsal spots aligned on the back of G. alba are larger and the fine spotting reduced which effectively enhances the darker dorsal pores in G. alba. This is one of three Western Australian frog species officially listed as ‘threatened and priority fauna’ and is considered to be ‘fauna that is rare or likely to become extinct’. It is one of the three southwest species with very small distributions that are the focus of a captive breeding project by the Perth Zoo. The other two species are G. vitellina and Spicospina flammocaerulea, also from dense swamps near the southern coast. Further Reading. Call analysis – Roberts et al. (1990); population biology – Driscoll (1997, 1998a, 1999a,b), Conroy and Brook (2003).

Genus GEOCRINIA

Geocrinia leai (Fletcher, 1898) Ticking Frog

! ! ! ! ! ! !! ! ! ! ! ! ! ! ! ! !! ! ! !! ! !!!! ! ! ! ! ! ! !!! ! ! ! ! ! !! ! ! !!! ! ! !! !!! ! ! ! ! ! !

Bridgetown (B. Maryan) Denmark (B. Maryan)

Description. A small brown or yellowish species with smooth ventral skin. The limbs are moderately long (TL/S-V 0.39–0.52). The toes are unusual in having the tips weakly dilated. There is a dark mid-dorsal patch usually with an irregular border; occasionally this patch is narrowly edged with white or the patch may be replaced by large, irregular yellow markings. Undersurface greenish. Males 13–22 mm; females 15–26 mm S-V. Distribution. Southwest region. Coastal and near coastal areas from Dandaragan south and east to Two Peoples Bay; inland to Chittering

Brook, Nannup and the Porongorups. Habitat. Associated with streams and swamps. Advertisement Call. A short high-pitched burst of notes, followed by a ‘tick…tick…tick’. Breeding Biology. A late autumn and winter breeder. Females lay clumps of eggs attached to vegetation above the waterline, and the larvae fall into the water below after they hatch. Tadpoles take more than 4 months to complete metamorphosis. Etymology. Named after A. M. Lea, a 19th Century Australian entomologist, who collected the specimens from which the species was described. Remarks. This is the only Western Australian species that lays its spawn attached to aquatic vegetation above the waterline, and the form of the toes also sets this species apart from other Geocrinia species. Further Reading. Tadpole description – Main (1957a).

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FROGS OF WESTERN AUSTRALIA

Geocrinia lutea (Main, 1963) Walpole Frog

! ! !

Walpole (B. Maryan)

Description. A relatively long-legged Geocrinia (TL/S-V 0.36–0.44), similar to G. rosea but lacking the pink belly. Skin smooth. The back is grey-brown with a darker mid-dorsal stripe (shaped somewhat like a molar tooth). There are pink spots in the groin and on the flanks and thighs. The belly is yellowish with scattered irregular dark spotting. Throat and undersurface of limbs are black in males. Males 18–23 mm; females 22 mm S-V.

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Distribution. Southwest region. Known only from a small area near Walpole and Nornalup, and north to Mount Frankland. Habitat. Wet forest litter on peaty sand. Advertisement Call. Similar to the ‘tk…tk…tk’ of G. rosea. Breeding Biology. Males call from tunnels in hummocks of mud covered with low matted vegetation. Egg masses of 25–30 are laid in small depressions. Larvae do not enter water but develop in the degraded jelly mass following hatching. Larvae collected at an early stage completed their development in 50 days. Etymology. lutea means ‘yellow’ in reference to the colour of the belly. Further Reading. Call analysis – Roberts and WardellJohnson (1995); population genetics – Driscoll (1998b); fire impact – Driscoll and Roberts (1997).

Genus GEOCRINIA

Geocrinia rosea (Harrison, 1927) Roseate Frog

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Pemberton (B. Maryan) Karri Valley (M.A. Cowan)

Description. A small, relatively short-legged species (TL/S-V 0.35–0.40) with smooth dorsal and ventral surfaces. The back is grey or brown with a broad, dark brown mid-dorsal stripe, often resembling a molar tooth. The undersurface of the body ranges from intense pink to red or fawn; the throat is dark slate or black in male frogs. Males 17–22 mm; females 24–25 mm S-V. Distribution. Southwest region. Donnelly River to Shannon and Gardiner River drainages in the southwest corner.

Habitat. Karri and Jarrah forests; individuals have been found beneath vegetation and rotten logs. Advertisement Call. A ‘tk...tk...tk’. Breeding Biology. The 25–30 eggs are laid from September to December out of water in shallow depressions besides streams, seepages and in upland sites in wet forests. The tadpole stages are passed in the degraded jelly mass following hatching. Emergent metamorphs have been observed in November. Etymology. rosea refers to the rose-coloured belly of this species. Further Reading. Call description – Main (1965a), Roberts and Wardell-Johnson (1995); life history and larvae – Main (1957a, 1965a), Watson and Martin (1973); population structure – Driscoll (1998b), Driscoll and Roberts (2007).

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FROGS OF WESTERN AUSTRALIA

Geocrinia vitellina Wardell-Johnson and Roberts, 1989 Orange or Yellow-bellied Frog

!

Spearwood Creek (B. Maryan)

Description. A small, muscular-limbed (TL/S-V 0.34–0.43) frog. Back brown to grey with darker spots, usually centred on pores which almost form flattened warts. Most prominent of these are two parallel rows extending from the eye to the cloaca and a vertebral row. Smaller brown spots are scattered evenly over the back. Ventral surface smooth and bright orange or the yellow of an egg yolk. Males 21–25 mm; female 18 mm S-V. Distribution. Southwest region. Only known from an area of about 4 km2 (mostly along Spearwood Creek) on the north bank of the

Blackwood River. It has the smallest known distribution of any Western Australian frog. Habitat. Most abundant at seepages on eastern slopes of Spearwood Creek, but rare on the floor of the valley. Advertisement Call. An irregularly repeated, discrete series of 9–15 pulses repeated slowly enough to be resolved by ear. Breeding Biology. A spring breeder, males call from September to December from small depressions under dense vegetation where the eggs are laid. Etymology. vittellina refers to the egg-yolk colour of the belly. Remarks. This is one of 3 Western Australian frog species listed as ‘threatened and priority fauna’ and is considered to be ‘fauna that is rare or likely to become extinct’. Further Reading. Call analysis and breeding biology – Roberts et al. (1990); population biology – Driscoll (1997, 1998a, 1999a,b), Conroy and Brook (2003).

Heleioporus eyrei, Emu Point (G. Harold) 52

Genus HELEIOPORUS Gray, 1841 Diversity and Evolutionary Relationships A small genus of robust large-bodied limnodynastine frogs. Recent genetic work indicates the sister taxon is the genus Limnodynastes (Frost et al. 2006), but other scenarios are possible (see Roberts and Watson 1993; Morgan et al. 2007). There are 6 species in southern Australia: 1 declining species in southeastern Australia (H. australiacus) with the remaining 5 species endemic to the southwest of Western Australia. Recent genetic work by Morgan et al. (2007) has provided a new perspective on relationships within Heleioporus. The eastern H. australiacus is the sister taxon to the 5 Western Australian species, and of the western species H. barycragus is the sister taxon to the remaining 4. Although H. eyrei and H. psammophilus are very similar morphologically and may even hybridise in some instances, H. psammophilus appears to be more closely related to H. albopunctatus.

Morphological Characteristics Head and body robust; head short; pupil vertical; tympanum not visible; upper jaw toothed; tongue broad; parotoid glands slightly raised. Limbs short; males with muscular arms. Fingers unwebbed; first finger longer than second and in most species males have large, thorn-like nuptial pads (Figure 21). Toes partly webbed; pale shovel-shaped inner metatarsal tubercle.

Reproductive Mode All species congregate to breed at the onset of autumn rains, with metamorphosis usually completed by late winter. Mixed breeding congregations involving 3 or more species occur in the Perth hills. Eggs are large and unpigmented and are laid singly in a foam mass at base of elongate breeding burrow dug into margin of seasonal swamp or bank of stream. Tadpoles hatch in the burrow and undergo initial development in the degraded jelly mass. When the reliable winter rains form ponds in early winter, the larvae are flushed from burrows and complete the rest of their development as a conventional aquatic tadpole. Lee (1967) provides a wealth of biological information and gives additional localities based on sight and sound records for all species of Heleioporus.

Etymology Not specifically stated by Gray, but probably from Greek helios (sun) and poros (holes), referring to the dorsal pattern of the first-named species, H. albopunctatus.

FROGS OF WESTERN AUSTRALIA

Heleioporus albopunctatus Gray, 1841 Western Spotted Frog

!! !! !! !! ! !!! ! !! !! ! ! ! !! !! ! ! ! ! ! !!! ! ! ! !! ! !! !!! !! !! ! !! ! ! !! ! !! !!!!! !! !!!! ! !! ! ! ! ! ! ! ! !! !! !! ! ! !! !! ! !! !!! !! ! ! ! !!! ! ! ! ! ! ! ! !! !!!! ! !!! ! !!! ! ! !! !!! !! !! ! !! ! !! !

!!

Boyagin Rock (B. Maryan)

Description. A large, robust, dark chocolatecoloured frog with short limbs (TL/S-V 0.29– 0.34). Adult males have 1 large sharp, black spine and 1–2 smaller spines on the first finger (Figure 21C). There are numerous, regularly spaced, large spots or blotches of white, cream or pale yellow over the entire back, sides and limbs. Males 56–79 mm; females 60–100 mm S-V. Distribution. Southwest region and adjacent arid zone. The wheatbelt and adjacent goldfields from Kalbarri National Park south to Tambellup and east to Jerramungup and Frank Hann National Park. Habitat. On clay and sandy soils throughout the southwest. Burrows are excavated near swamps, on the banks (or under stones) of shallow, ephemeral watercourses and in the steep edges of claypans. Advertisement Call. A short, high-pitched ‘coo’

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that is repeated at a rate of slightly more than once per second. Breeding Biology. Females lay 250–700 eggs in a burrow in autumn in a foam nest, and take from 10–30 days to hatch; metamorphosis is in September and October. Tadpoles have been found in colloidal water or in clear ponds. The dorsal surfaces are iridescent gold with scattered, minute, black spots; there are 6–7 rows of upper and 3 rows of lower labial teeth. Etymology. albo means white and punctatus means spot; thus ‘white-spotted’. Further Reading. Evolutionary relationships – Morgan et al. (2007); population biology – Davis and Roberts (2005a); call analysis – Littlejohn and Main (1959); breeding biology – Lee (1967), Davis and Roberts (2005b); tadpoles – Davies (1991); physiology – Withers (1993); egg parasite – Davis and Disney (2003).

Genus HELEIOPORUS

Heleioporus barycragus Lee, 1967 Hooting Frog

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Darlington (B. Maryan)

Description. A large and robust species, with a stocky appearance, short legs (TL/S-V 0.28– 0.35) and large protruding eyes. It is coloured light rich chocolate with numerous small bright yellow spots or blotches on the flanks. In some individuals there are small black spines at the center of each of the yellow spots on the flanks. Males have a large, black, thorn-like, nuptial spine on the first finger, and other smaller spines on the first and second fingers (Figure 21D). Males 60–83 mm; females 62–86 mm S-V. Distribution. Southwest region. From Bullsbrook south to Darkan, east to Northam and including Dryandra National Park and Wandoo forest. Habitat. Occurs in rocky forests and breeds

along flowing watercourses on clay or granite. Advertisement Call. Like the slow hooting of an owl. Breeding Biology. Eggs are laid in autumn in a foam nest in a relatively shallow burrow that is excavated by the male in the side of a vertical bank. The tadpole grows to 60 mm in length and is mottled with black or brown above and usually has a lighter mid-vertebral stripe. Etymology. barycragus means ‘deep-voiced’. Further Reading. Evolutionary relationships – Morgan et al. (2007); call analysis (as H. australiacus) – Littlejohn and Main (1959); reproduction and tadpoles – Lee (1967).

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FROGS OF WESTERN AUSTRALIA

Heleioporus eyrei (Gray, 1845) Moaning Frog

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Perry Lakes (M. Peterson)

Description. A robust medium-large frog with short legs (TL/S-V 0.30–0.33). Back brown or dark slate grey, largely obscured with irregularly defined patches of dull yellow or pale grey. Parotoid gland area usually with a yellowy wash. Adult males are without the typical black nuptial spines on the first or second fingers of other Heleioporus species (Figure 21A). Males 33–66 mm; females 46–63 mm S-V. Distribution. Southwest region. Coastal and near coastal districts from Geraldton south and east to Cape Arid. Inland to Marchagee, Corrigin, Jerramungup and Gibson Soak. Also on Rottnest and Bald Islands. Habitat. Found in swamps with sandy soil. Advertisement Call. A long and rising low moan. Breeding Biology. Calling begins in autumn after the first heavy rains of the year in areas

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that will flood later in winter. Clutches of 80– 500 eggs are laid in a burrow dug at an angle to a horizontal land surface. Larvae hatch and remain in the foamy egg mass until the burrow is flooded. Tadpoles feed for several more months before metamorphosis in September or October. They are densely mottled with black and gold, have a red or gold vertebral stripe and a curved lateral line. Etymology. Named for Edward John Eyre who, with his Aboriginal guide Wylie, was the first white person to walk across the Nullarbor Plain in 1841. The species was described in an appendix to Eyre’s account of his travels. Further Reading. Call analysis – Littlejohn and Main (1959); reproduction – Dakin (1920), Packer (1963), Lee (1967); tadpoles – Packer (1966), Lee (1967); physiology – Packer (1963), Lee (1968).

Genus HELEIOPORUS

Heleioporus inornatus Lee and Main, 1954 Whooping Frog

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Brookton Highway (B. Maryan)

Description. A large robust frog with protruding eyes and moderately short legs (TL/S-V 0.31– 36). Back and sides uniform coppery brown without markings, or brown faintly mottled with pale yellow, grey or white. Adult male with 1–2 small black nuptial spines on the first finger (Figure 21B). Males 44–64 mm; females 57–73 mm S-V. Distribution. Southwest region. Darling Range from Chidlow south to the vicinity of Margaret River, and along the south coast east almost to Albany. Habitat. Prefers sandy or peaty soil. Advertisement Call. A loud ‘ whoop…whoop… whoop’ repeated frequently.

Breeding Biology. Breeds in autumn before the winter rains. Approximately 100–250 large, pale yellow eggs are laid in autumn in a foam nest constructed in a 20–40 cm deep burrow often penetrating the side of sloping ground, or in the side of a bank. Tadpoles have long tails and narrow fins and are very dark with a reddishbrown or yellow vertebral stripe. Metamorphosis occurs in spring. Etymology. inornata means ‘inornate’ or unadorned relative to other Heleioporus. Further Reading. Call analysis – Littlejohn and Main (1959); reproduction and tadpoles – Lee (1967).

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FROGS OF WESTERN AUSTRALIA

Heleioporus psammophilus Lee and Main, 1954 Sand Frog

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!! ! ! ! !

The Lakes (B. Maryan)

Description. A robust species with short limbs (TL/S-V 0.28–0.35). Back brown or grey with marbling of grey, white, dull yellow or paler brown. Sides dotted with white. In northern populations, the adult male has a black nuptial spine on the first finger (Figure 21B) but it is usually absent in the southern populations, making H. psammophilus more difficult to distinguish from H. eyrei. Males 33–62 mm; females 41–60 mm S-V. Distribution. Southwest region. On the west coast, north to the Irwin River south to Bussleton and inland to Coorow and Dongolocking. On the south coast from Windy Harbour east to Jerramungup. Habitat. Angular fine-grained sands and sandy clays.

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Advertisement Call. A short high-pitched note repeated frequently. More high-pitched and faster than any other Heleioporus. Breeding Biology. An autumn breeder. Males call from burrows dug into sandy soils that will later flood in winter. The tadpoles of this species are difficult to distinguish from the closelyrelated H. eyrei. Etymology. psammophilus means ‘sand-loving’, in reference to this species’ preference for sandy soils. Further Reading. Call analysis – Littlejohn and Main (1959); reproduction and tadpoles – Lee (1967); population genetics – Berry (2001).

Genus HELEIOPORUS

a

B

C

D

Figure 21

Variation in nuptial pads of Heleioporus species: A) H. eyrei and southern H. psammophilus (no pad); B) H. inornatus and northern H. psammophilus; C) H. albopunctatus; D) H. barycragus. Ventral surface shown in (AC) and the dorsal surface in (D).

Heleioporus albopunctatus burrow entrance (M. Cowan)

Heleioporus albopunctatus embryos in foam nest (BB Wells / DEC) 59

Genus LIMNODYNASTES Fitzinger, 1843 Diversity and Evolutionary Relationships A moderately large genus of limnodynastine frogs, with a total of 11 species. Heleioporus may be the sister genus (Frost et al. 2006). Represented in Australia in the mesic southeast and southwest and the tropical north, extending into southern New Guinea. In Western Australia, 1 species is endemic to the southwest region and 3 occur in the Kimberley. Limnodynastes has experienced some key changes in content during this decade. Genetic work by Schäuble et al. (2000) and Frost et al. (2006) indicate that Megistolotis Tyler, Martin and Davies, 1979 is embedded within Limnodynastes, despite having striking differences from other Limnodynastes (e.g. large distinct tympanum and rock-dwelling habits). Furthermore, L. ornatus and L. spenceri are not closely related to other Limnodynastes, instead having stronger affinities to Lechriodus from eastern Australia and New Guinea. These species were transferred to Opisthodon Steindachner, 1867 by Frost et al. (2006), but are allocated to Platyplectrum Günther, 1863 (the older available name) in this volume. The similarsounding Platyplectron Peters, 1863 was erected for L. dumerilii in the east which was synonomized into Limnodynastes by Neiden (1923) which may have led to the confusion.

Morphological Characteristics Body robust or flattened, head large; pupil vertical; tympanum visible but with indistinct annulus (Figure 3B), except L. lignarius with prominent annulus; upper jaw toothed; vomerine teeth behind internal opening of nostrils; tongue broad. Skin glands present or absent. Limbs moderately long. Fingers unwebbed; toes webbed or unwebbed.

Reproductive Mode Aquatic reproduction. Eggs pigmented. All species produce a foam nest in which the eggs are placed as they are fertilized. The females have specialized flanges on the fingers which are used to trap bubbles for the construction of the nest. Tadpoles wriggle through the degraded nest to complete development in water as typical anuran larvae.

Etymology Limnos means marsh, lake or pool, and dynastes refers to a dynastic ruler; usually interpreted as ‘Lord of the marsh’.

Further Reading Evolutionary relationships – Roberts and Maxon (1986), Schäuble et al. (2000), Frost et al. (2006); foam nests – Tyler and Davies (1979a), Tyler et al. (1983a).

Limnodynastes lignarius, Kakadu, NT (G. Harold) 61

FROGS OF WESTERN AUSTRALIA

Limnodynastes convexiusculus (Macleay, 1877) Marbled Frog

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! !! !! ! ! !

King Leopold Ranges (G. Gaikhorst)

Description. A squat-bodied frog. The limbs are of moderate length and build (TL/S-V 0.36–0.42). The fingers and toes are long and unwebbed. The outer metatarsal tubercle is moderately developed (similar in size to fifth toe). Most individuals are pale olive with numerous small patches of dark green arranged in a regular pattern. The dark green patches may bear a narrow border of cream or deep crimson, and there is a prominent pale yellow or cream labial gland at the angle of the jaw. Quite commonly the dark patches are raised above the surface and in large individuals there may be tiny blacktipped spines crowning the raised areas. Males 39–58 mm; females 46–61 mm S-V. Distribution. Kimberley region. South to Beverley Springs, Mount Barnett and Kununurra.

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Also widely distributed in the northern portion of the Northern Territory, northeast Queensland and southern New Guinea. Habitat. Swamps, grasslands and other lowlying areas exposed to seasonal flooding. Advertisement Call. A loud repeated honking. The male calls from water at the base of grass, sometimes occupying holes of the crab genus Holthuisiana. Breeding Biology. A wet season breeder, with males calling from December to March, and even during periods of no rain. Tadpoles are completely black and may reach 60–70 mm length. Etymology. convexiusculus means ‘arched rump’ in reference to the posture often displayed by this species.

Genus LIMNODYNASTES

Limnodynastes depressus Tyler, 1976 Flat-headed Frog

! !!! !!! ! !

♂ Kununurra (P. Doughty) ♁ Kununurra (P. Doughty)

Description. A medium-sized, squat species with moderate limbs (TL/S-V 0.36–0.44), flattened head and widely separated eyes with prominent upper eyelids. Males possess massive arms with a protruding terminal phalanx on the fourth finger. The hands and feet are unwebbed. Outer metatarsal tubercle only weakly developed. The back and head are pale brown and covered with numerous large spots of dark brown, with white undersurfaces. Males 31–54 mm; females 34–50 mm S-V. Distribution. Kimberley region. Occuring around Kununurra and Lissadell Station. They extend in the Northern Territory from the Keep River to the Bradshaw Country. Habitat. Swamps and flooded grasslands.

Advertisement Call. A rapid series of about 7 notes repeated rapidly. Calling site is the edge of shallow water or from crab burrows. Breeding Biology. Eggs are laid in a small foam nest (8 x 5 cm) within or above a depression in the ground. Spawn can reach temperatures of 41°C. Tadpoles attain a maximum length of 80 mm. Etymology. depressus means ‘flat’, in reference to the flattened appearance of the head of this species. Remarks. Identification of L. depressus in the eastern Kimberley has been difficult in the past, owing to confusion with presumably introduced L. tasmaniensis from South Australia and the belief that the species was extinct when the only known site of occurrence was destroyed by Lake Argyle. Fortunately, neither scenario is true, and populations occur around Kununurra and Wyndham and extend eastwards into the Bradshaw Country of the Northern Territory. Further Reading. Anatomical description – Davies and Burton (2000); confusion with L. tasmaniensis – Martin and Tyler (1978); developmental biology – Tyler and Davies (2000).

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FROGS OF WESTERN AUSTRALIA

Limnodynastes dorsalis (Gray, 1841) Western Banjo Frog

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Three Springs (G. Harold) (L) Cape Naturaliste (B. Mayran) (R) Foam nest construction (M. Cowan)

Description. A robust species with a large triangular head and moderately short legs (TL/S-V 0.34–0.45). Distinguished from other Western Australian species by the presence of a large protruding oval gland on the upper surface of the calf. The toes are very slightly webbed and the foot bears a large shovel-shaped, inner metatarsal tubercle (Figure 9). The back varies from pale brown to dark chocolate bearing extensive areas of deep green or olive, and usually a very narrow cream or pale yellow mid-dorsal stripe. There is a prominent golden supralabial gland (Figure 6B) and the groin is entirely crimson. Males 28–64 mm; females 54–73 mm S-V. Distribution. Southwest region and adjacent arid zone. The Lower Murchison River south and east to almost Wattle Camp. Inland to Galena, Moorine Rock, Lake Cronin, Peak Charles and Coragina Rock. Habitat. Found in a variety of habitats, including 64

forests, swamps, grasslands and the wheatbelt. During the winter months it is found among vegetation near permanent water. A burrowing species that submerges itself during dry conditions. Advertisement Call. A deep and explosive ‘bonk’, repeated slowly, like the plucking of a banjo string. Males call from the shelter of dense vegetation or other sites where they can remain hidden. Breeding Biology. Spawn laid in winter and spring in a large foam nest on the surface of static or slowly flowing water. Tadpoles (Figures 17A and 18A) are large (to 65 mm), black and with deep tail fins. Etymology. Like most names coined by Gray, he did not specify what the name meant, but dorsalis refers to the back, and may refer to the dorsal pattern of the holotype specimen. Further Reading. Call analysis – Martin (1972); breeding biology and tadpoles – Main (1965a).

Genus LIMNODYNASTES

Limnodynastes lignarius (Tyler, Martin and Davies, 1979) Woodworker or Carpenter Frog

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!

! !!!! !

!!! ! !

Mornington Station (P. Doughty)

Description. A medium-sized frog, with head and body broad and flattened. Characterised by an exceptionally large and conspicuous tympanum. Fingers and toes are elongate, with a trace of basal webbing between the toes. The limbs are moderately short (TL/S-V 0.36–0.45). The inner metatarsal tubercle is not modified for digging. Males have enlarged forearm muscles, and in the breeding season there is a small cluster of black, thorn-like spikes on the first finger, and small spike-tipped tubercles over the entire dorsal surface. The back is an irregularly marked dull reddish slate or dark olive, occasionally flecked with silver or gold. Males 43–62 mm; females 47–61 mm S-V. Distribution. Kimberley region. South to Mount Hart and Lake Argyle. Extends to the Arnhem Land escarpment of the Northern Territory. Habitat. Always associated with rocks. Occurs beneath rock piles or scree slopes, open escarpments, caves or among boulders in shallow slow-flowing streams.

Advertisement Call. A single, soft tap resembling a piece of timber being struck with a hammer. The call is repeated at regular intervals of roughly once per second. Breeding Biology. A tropical wet season breeder. Females lay 300–400 pigmented eggs in a foam nest in rock pools, or beneath rocks at the edge of water-draining scree slopes. The tadpoles become an intense black colour and have a flattened appearance with vertral sucker-like mouths. They reach a maximum length of 55 mm and may complete their development in less than 9 weeks. Etymology. lignarius means ‘carpenter’ or ‘woodworker’. Further Reading. Call analysis and tadpoles – Tyler et al. (1979); evolutionary relationships – Hutchinson and Maxson (1987a), Schäuble et al. (2000), Frost et al. (2006).

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Genus LITORIA Tschudi, 1838 Diversity and Evolutionary Relationships A large and complex genus of hylid frogs, with 68 species in Australia. Also well-represented in New Guinea and extending to neighboring parts of Indonesia, Timor and the Solomon Islands. Introduced to Guam, Vanuatu, New Zealand and the United Kingdom. A total of 19 species occur in Western Australia, with 16 in the Kimberley, 3 in the southwest and 1 in the arid zone. The Litoria aurea species-group (including the southwest region’s L. cyclorhyncha and L. moorei ) is more closely related to the burrowing hylids of the genus Cyclorana than with other Litoria species (Tyler 1978, 1979). This view is supported by morphological (Burton 1996), genetic (Frost et al. 2006) and biochemical (Maxson et al. 1985; Hutchinson and Maxson 1986, 1987b) analyses. The aquatic L. dahlii is also closely related but its placement is more problematic (Burns and Crayn 2006). The combined evidence suggests the following overview of Western Australian Litoria. The southwest endemic L. adelaidensis appears to be a basal member with no close relatives. The other two southwestern species L. moorei and L. cyclorhyncha are closely related to each other and show clear links to members of the L. aurea species-group of southeastern Australia. Recent genetic work (Burns and Crayn 2006) indicate that the two southwest taxa are likely of subspecific rank, differing genetically by less than 1%. Should these two taxa be synonomized, L. cyclorhyncha is the older available name. Litoria rubella of the arid zone and Kimberley is very distinct in the Australian context, but has possible links to various New Guinean species (Tyler and Davies 1978; Menzies 2006). Litoria caerulea, L. cavernicola and L. splendida from the Kimberley form a distinctive group of large tree frogs in the genus; the generic name Pelodryas Günther, 1859 will likely apply to this group of species. The three other arboreal species are L. bicolor, L. microbelos and L. rothii, with each being quite distinctive from the others. The immediate affinities of L. microbelos are probably with several New Guinean species (Tyler and Davies 1978). The ‘L. freycineti assemblage’ of ground frogs (including the type species of Litoria) is well represented in the Kimberley by L. inermis, L. nasuta, L. pallida, L. tornieri and L. watjulumensis. The rock-dwelling L. coplandi and L. staccato appear to be closely related, with the smaller-bodied L. meiriana only distantly related.

Morphological Characteristics A highly variable group of species owing to diverse ecologies. Head and body generally elongated. Pupil horizontal, with or without ventral extension; tympanum at least partly visible; upper jaw toothed; vomerine teeth present or absent; tongue long or broad. Skin glands visible or not visible. Limbs short, moderately long or long; fingers and toes with dilated terminal discs (but minimal in L. dahlii, L. inermis, L. nasuta, L. pallida and L. tornieri ); fingers webbed or unwebbed; toes webbed.

Reproductive Mode Aquatic reproduction. Eggs pigmented; spawn laid as single eggs, small groups or clumps on or beneath surface of water. Tadpoles of all species develop in water but show a great variety of morphological adaptations (Anstis 2002).

Etymology Litoria may be derived from ‘shore’, ‘strand’ or ‘beach’; alternatively, the name may have been coined merely for the way it sounds as Tschudi did not have knowledge of the habitat of the first-described species, L. freycineti.

Further Reading Tyler and Davies (1978, 1993), Roberts and Watson (1993), Tyler (1994), Plötner et al. (2007).

Litoria caerulea, Lakeland, QLD (G. Harold)

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FROGS OF WESTERN AUSTRALIA

Litoria adelaidensis (Gray, 1841) Slender Tree Frog

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South Perth (B. Maryan) Margaret River (B. Maryan)

Description. An elongate frog with a narrow, tapering head and slender body. The fingers are unwebbed and elongate with distinctly expanded discs. The legs are moderately long (TL/S-V 0.47–0.52) and the toes are extensively webbed. The dorsal colour ranges from unmarked brown or green to brown with large green patches. There is usually a broad, dark brown stripe along the side of the head and body; from the eye to the flanks this stripe is bounded by a narrow, brilliant white stripe. There are brilliant small red or orange spots on the back of the thigh. Males 34–45 mm; females 42–47 mm S-V. Distribution. Southwest region. North from Port Gregory south and east to Bremer Bay and from Esperance to Mount Ragged. Inland to Mundaring Weir, Dwellingup and Harvey on

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the west coast and Manjimup to Tambellup on the south coast. Habitat. Dense vegetation at the edge of static or slowly-moving water. A common inhabitant of backyard ponds in urban areas. Advertisement Call. A grinding sound or ‘harsh grating screech’ (Main 1965a). Males call from water, adjacent soil, or from vegetation as much as 1 m above the ground. Breeding Biology. This species usually breeds in early spring. Calls, however, can be heard from autumn to mid-summer. The spawn is laid in an irregular mass attached to vegetation just below the surface of the water. Tadpoles have 2 upper and 3 lower rows of labial teeth and high, deep fins. Mature tadpoles are dark brown with a pair of pale stripes extending backwards from the nostrils. Metamorphosis occurs in late spring or early summer. Etymology. Named for Adelaide, South Australia (where L. adelaidensis does not occur), erroneously believed to be the location of collection by Gray, who worked in London and never visited Australia. Further Reading. General – Main (1965a); call analysis – Smith and Roberts (2003d).

Genus LITORIA

Litoria bicolor (Gray, 1842) Northern Dwarf Tree Frog

! !

!! ! !! ! ! !! ! !! !! ! !! !! ! ! !!

!

Mornington Station (J. Francis)

Description. A small, slender and rather delicate climbing frog with a flattened narrow, triangular head and elongated body. Limbs are moderately long (TL/S-V 0.47–0.51). The fingers are short, with small discs and a trace of web at the base. The toes are fully webbed. Most adults are pale green but some are a dull bronze. Juveniles are bronze or brown and may gradually acquire green colouration when about three-quarters of adult size. There is a narrow brown stripe on the side of the head and body, bounded beneath by a brilliant white stripe. The hidden portions of the limbs are orange or bluish purple. Males 23–30 mm; females 25–35 mm S-V. Distribution. Kimberley region. South to Yampi Peninsula, Beverley Springs and the Lower Ord. Also on Sir Graham Moore Island. They extend eastwards across tropical Northern Territory. Habitat. Most abundant in areas with dense grass and reeds such as swamps.

Advertisement Call. A high penetrating call – ‘reek…reek…reek’, resembling a wooden ratchet. Males commonly call when in a vertical position clinging to grass stems in flooded areas. However, they can call in vegetation as much as 5 m above the ground. Breeding Biology. The eggs are small (1 mm in diameter), pigmented and laid in small clumps of 10–20 attached to submerged vegetation. The tadpoles are a brilliant pale brown and exhibit a white stripe beneath the eye in the latter stages of development. They attain a maximum length of 40 mm. Etymology. bicolor means ‘two colours’, referring to the contrast between the flanks and belly. Further Reading. Evolutionary relationships – Tyler and Davies (1978); Menzies et al. (2008).

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FROGS OF WESTERN AUSTRALIA

Litoria caerulea (White, 1790) Green Tree Frog

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Kununurra (B. Maryan)

Description. A large species with a medium-large build. Parotoid glands and supratympanic folds are prominent and partly cover the tympanum. The fingers (Figure 11B) have very broad fringes and are partly webbed while the toes are webbed extensively (Figure 10A). The limbs are of moderate length (TL/S-V 0.44–0.50). Bright green or olive, occasionally lightly marked with white (mainly on the flanks). The hidden portions of the limbs and flanks are commonly pale yellow. Males 67–77 mm; females 60–110 mm S-V. Distribution. Kimberley region. South to Anna Plains, Edgar Range, Noonkanbah, Fitzroy Crossing and the Elvirie River. Elsewhere it extends east to Queensland and south to South Australia and New South Wales. Also scattered locations in southern New Guinea. Habitat. Shelters in tree hollows and rock crevices. One of the most common frogs associated with human-made structures. Advertisement Call. A deep barking sound repeated regularly; sometimes resembles timber being sawn or the barking of large dogs. Males will call in unison during the day in response to showers.

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Breeding Biology. Breeding takes place between November and March. Several thousand (1500– 4400) small, pigmented eggs are laid upon the surface of the water. The spawn sinks within 24 hours. Tadpoles attain a maximum length of 63 mm. Tadpole growth is rapid and metamorphosis is completed in 4–5 weeks. Etymology. caerulea means ‘blue’, the colour of specimens preserved in alcohol which dissolves the yellow pigment in the skin. It is possible that the names in White’s journal were coined by George Shaw, John Hunter or his editor Thomas Wilson, who would have seen only preserved specimens. The short description is most likely the work of Shaw. Remarks. One of the best known of all Australian species, they are amenable to being handled and are very popular as pets, both in Australia and overseas. One breeder in the USA breeds over 60,000 per year! Further Reading. Breeding biology – Anstis (2002); as pets – Tyler (1996).

Genus LITORIA

Litoria cavernicola Tyler and Davies, 1979 Cave Frog

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Mitchell Plateau (G. Gaikhorst)

Description. A moderate-sized species with a flattened appearance and possessing mediumlong hindlimbs (TL/S-V 0.42–0.50). There is a large and distinct tympanum and conspicuously expanded discs on the fingers and toes; the toes are extensively webbed. Green to reddish-brown with coarsely granular dorsal skin. Males 45–60 mm; females 50–57 mm S-V. Distribution. Kimberley region. From the Mitchell Plateau south and west to the Prince Regent River Nature Reserve. Also on Bigge Island and Katers Island.

Habitat. Occurs in caves, ledges or trees on massive sandstone boulders and rock platforms. Advertisement Call. A soft repetitive honking. Breeding Biology. Males call from trees among large boulders or caves, or by shallow pools on rock platforms. Eggs are laid in shallow rock pools. Etymology. cavernicola means ‘cave-dwelling’.

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FROGS OF WESTERN AUSTRALIA ! ! !! ! ! ! ! ! ! ! ! ! ! ! ! !! !! ! ! ! ! !! !!! ! ! ! ! ! ! ! ! ! !!!! ! !! ! ! ! ! !! ! !!! ! !! ! !! !! ! ! ! ! ! ! ! ! ! !! ! ! ! ! ! ! ! !!

Litoria coplandi (Tyler, 1968) Rock Frog

Wyndham (P. Doughty)

Description. A small to moderate-sized species. The tympanum is large and conspicuous. The fingers are long, slender, unwebbed and unfringed and have conspicuous expanded terminal discs. The hindlegs are moderately long to long (TL/ S-V 0.45–0.65) with the toes fully webbed and the terminal disks moderately expanded. Usually a plain, pale brown. The top of the head from the tip of the snout to a position between the eyes is lighter in colour, with a transocular bar visible in most individuals. Males 17–36 mm; females 39–42 mm S-V. Distribution. Kimberley region. South to Mount Anderson in the west and Denison Range in the east. Also found in tropical Northern Territory and extending into Queensland. Habitat. Rock-dwelling; common along rocky creeks and waterfalls in the wet season. Advertisement Call. A soft repetitive whirring that increases in amplitude, interspersed with intermittent short calls.

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Breeding Biology. An early wet season breeder. Eggs are scattered singly, falling to the floor of rock pools or slow-moving temporary streams in rocky areas. The tadpole is black and has a conspicuous, large, white spot on each side of the body. Most tadpoles attain a total length of more than 45 mm. Etymology. Named for Stephen J. Copland for contributions to Australian frog taxonomy, including a revision of Litoria (as Hyla) in 1957. Remarks. This species is exceedingly agile and therefore difficult to capture. Although most likely to be seen at the edge of water, it is capable of climbing, particularly upon quite smooth cliff faces. Further Reading. Call description – Doughty and Anstis (2007).

Genus LITORIA

Litoria cyclorhyncha (Boulenger, 1882) Spotted-thighed Frog

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Lake Cronin (G. Gaikhorst)

Description. An elongated, moderately large to large-sized frog with long limbs (TL/S-V 0.43–0.55). The fingers are unwebbed and bear prominent discs (Figure 11C); there is extensive webbing between the toes with moderately wide discs. The back is variable in colour, from a dark green to brown dappled with small, gold or bronze patches. The groin and the back of the thighs bear numerous clear, pale-yellow patches. Males 56–66 mm; females 63–80 mm S-V. Distribution. Southwest region and adjacent arid zone in the eastern portion of its range. From near Albany (where an ill-defined hybrid zone with L. moorei exists), inland to Broomehill, Lake Varley and Ravensthorpe. Also from Dalyup River east to Israelite Bay and inland to Scaddan and Coragina Rock. Occurs on Middle Island of the Recherche Archipelago. Habitat. Occurs near swamps and permanent water.

a

Advertisement Call. A low rising pitch followed by a series of growls. Breeding Biology. Details have not been reported, but presumably similar to L. moorei which has been more extensively studied. Calls have been heard in spring and summer. Etymology. cyclo means ‘circle’ and rhyncha means ‘snout’ in reference to this species’ more rounded snout relative to L. aurea which it was distinguished from. Remarks. Very closely related to L. moorei to the west. Indeed, recent genetic evidence indicates that L. cyclorhyncha may be the same species as L. moorei, or at best subspecies (Burns and Crayn 2006). However, further work is necessary to resolve the issue. Further Reading. Evolutionary relationships – Burns and Crayn (2006); reproduction – Main (1965a).

B

Figure 22

Patterning on inside of hind limbs: a) Litoria cyclorhyncha and b) L. moorei. 73

FROGS OF WESTERN AUSTRALIA

Litoria dahlii (Boulenger, 1896) Aquatic Frog

!

Humpty Doo, NT (B. Maryan)

Description. A moderately large to large and highly active species with a rather flattened body and moderately long limbs (TL/S-V 0.45–0.51). The fingers are long, pointed and unwebbed; the toes are pointed and extensively webbed. The dorsal surface is highly variable in colour, ranging from dark slate to a pale and rather diffuse green. There is usually a pale mid-vertebral stripe, and the back of the thighs bear white or grey spots. Males 49–63 mm; females 59–71 mm S-V. Distribution. Kimberley region. Known from near Wyndham in Western Australia, but extends east through tropical Northern Territory and Queensland and enters southern New Guinea. Habitat. Swamps and billabongs, where it is a top predator of other frog species (and often it’s own species). Advertisement Call. Unknown; reports of soft calls made while floating or even underwater. Breeding Biology. Observations on the sequence of breeding of frogs in the Northern Territory indicate that L. dahlii is probably one of the last

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species to breed in each wet season (February to March). The tadpole is distinctive, having broad longitudinal stripes along the tail. Tadpoles attain a maximum length of nearly 60 mm. Etymology. Named after Knut Dahl (1871– 1951) who led a Norwegian expedition in northern Australia from 1893–1896 which led to the collection of the specimens used for the description of this species. Remarks. In comparison with other Western Australian frogs this is an exceptionally timid creature. In the daytime it commonly basks on the ground beside water. In Western Australia, a single individual was collected 40 years ago with no reports since then, but probably occurs close to the Northern Territory border in blacksoil plains which are impassable in the wet season. Further Reading. Evolutionary relationships – Tyler et al. (1978), Burns and Crayn (2006).

Genus LITORIA

Litoria inermis (Peters, 1867) Bumpy Rocket Frog

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Leopold Ranges, (G. Gaikhorst)

Description. A small ground-dwelling frog with a prominent elongate head, long unwebbed fingers and moderately long to long hindlegs (TL/S-V 0.52–0.60) and incompletely webbed toes. The skin on the back is coarsely tubercular. The back is generally mottled with irregular patches of dark brown on a lighter background. Markings on the thighs are diffuse. There is only a diffuse loreal stripe with a distinct white patch at the angle of the jaw. Males 24–34 mm; females 29–35 mm S-V. Distribution. Kimberley region. South to Meeda, Mount Percy, Mount Barnett and the Denison Range. Also occurs in tropical Northern Territory and Queensland. Habtitat. Grasslands and streamsides. Advertisement Call. Complex: a loud constant computer-like ‘meep…meep’ repeated frequently,

followed by similar notes except with a waver; intermittent short soft calls are given between the longer louder series. Breeding Biology. Males call from open areas near water during the wet season. Spawn is laid in temporary pools where there is little or no emergent vegetation. Etymology. inermis means ‘unarmed’ in reference to the lack of a strong metatarsal tubercle for digging. Remarks. Very similar to L. pallida, but L. inermis is smaller with a blunter snout, has rougher skin, lacks a clearly-defined head stripe and the call involves constant and wavering main notes. Further Reading. Redescription, call analysis, reproduction and tadpoles – Davies et al. (1983); generic status and radiographs of feet – Strahan (1969).

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FROGS OF WESTERN AUSTRALIA

Litoria meiriana (Tyler, 1969) Rockhole Frog

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! !! !! !! !

!!

Kununurra (J. Francis)

Description. A very small and active frog with moderately long to long legs (TL/S-V 0.48– 0.56). The fingers are short, unwebbed and bear distinct expanded discs. The toes are fully webbed and also have pronounced circular discs. The dorsal surface is mottled slate or brown and there are commonly very small creamish or yellow spots or blotches on the backs of the thighs. Males 16–22 mm; females 18–23 mm S-V. Distribution. Kimberley region. South to Napier Range, Fossil Downs, Carson River, Dunham River Station and Kununurra. Also extends across the northern portion of the Northern Territory as far as Arnhem Land. Habitat. Found near rock pools and temporary creeks draining rocky areas. Also occurs in caves. Advertisement Call. The main call is a series of short high-pitched penetrating notes, with sharp

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notes given intermittently between the main calls. Breeding Biology. Eggs are laid in small static pools on or at the bottom of rock escarpments. One clutch included 39 eggs. The eggs are laid in a film upon the submerged surface of the rock and development takes approximately 3 weeks. Metamorphs are very small (8–10 mm S-V). Etymology. meiriana is the local Aboriginal name near the Northern Territory type locality for ‘rockhole’, which this species frequents. Remarks. For much of the year this species is diurnal as well as nocturnal. When disturbed it can skip across the surface of the water to escape, and will clamber up nearly vertical rock surfaces making it extremely difficult to capture. Further Reading. Call description – Doughty and Anstis (2007); development – Tyler and Crook (1987).

Genus LITORIA

Litoria microbelos (Cogger, 1966) Javelin Frog

! ! !

!

Drysdale River NP (P. Doughty)

Description. An exceptionally small, pale brown species with an elongated and sharply pointed head and moderately long to long slender limbs (TL/S-V 0.50–0.58). The fingers are unwebbed and bear small terminal discs. The toes are very long with reduced webbing, leaving at least the terminal two segments unwebbed. It has a darker brown stripe on the sides of the head and body, surmounting a narrow white stripe commencing beneath the eye and passing backwards, often as far as the groin. Males 14–16 mm; females 15–17 mm S-V. Distribution. Kimberley region. Only known from 2 sites; on the Mitchell Plateau and northern Drysdale River National Park. Also occurring across the peripheral northern coastline of Australia. Habitat. Found in dense tussocks of lowgrowing grasses, and elsewhere has been found to hide during the daytime beneath large rocks and logs.

Advertisement Call. The call is a high-pitched, soft buzz. Males call from vegetation rarely more than 30 cm from the ground. Breeding Biology. Breeding has been observed in January and February. Spawn clumps of approximately 60–275 eggs are laid in shallow water in temporarily flooded areas. The ova are less than 1 mm in diameter. Etymology. microbelos means small spear or javelin, alluding to the slender pointed body and head shape. Remarks. Litoria microbelos is the smallest member of the family Hylidae in Australia. When disturbed it is very active and difficult to capture. Further Reading. Reproduction and tadpoles – Anstis and Tyler (2005).

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FROGS OF WESTERN AUSTRALIA

Litoria moorei (Copland, 1957) Motorbike Frog

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Pelican Island (B. Maryan)

Description. A robust, muscular frog with a triangular-shaped head and moderately long (TL/S-V 0.46–0.58) and powerful hindlimbs. The fingers are long and unwebbed and have small but distinct discs on the tips. The toes are long with a moderate extent of webbing between them. The skin of the back generally bears distinct, isolated tubercles. The colouration is variable and individuals are capable of striking colour changes. The back ranges from uniform pale brown or green (usually with a paler middorsal stripe) to very pale brown with numerous, clearly defined very dark patches. There is a dark stripe from the tip of the nose to the eye, and behind the eye, through the tympanum to above the insertion of the forearm. The groin and back of the thighs is pale green and lacks the bold, yellow patches seen in the closely-related L. cyclorhyncha (Figure 22). Males 47–71 mm; females 53–78 mm S-V. Distribution. Southwest region. From the Murchison River south and east to Pallinup River, inland to Three Springs, Dalwallinu and Lake Dumbleyung. A poorly-defined hybrid zone occurs from near Albany and to the north. A small population persists on Rottnest Island. Habitat. Areas of permanent water, often climbing to hide beneath bark or else found beneath rocks or logs. A common resident of 78

backyard ponds in Perth and other southwestern towns. Advertisement Call. A series of rising notes that resemble the sound of a motorbike changing gears, followed by a series of low growls. Breeding Biology. Eggs are laid in spring to mid-summer. The spawn clump is a large floating mass attached to vegetation. Larvae can be found throughout summer and often metamorphosis is as late as April, although some individuals metamorphose the following spring. Tadpoles have deep fins, a pointed tail-tip and are very dark. Etymology. Named after John A. Moore (1915– 2002), an American herpetologist who was a pioneer of Australian frog studies in the 1950s. Remarks. Recent work indicates very little genetic difference between L. cyclorhyncha and L. moorei (Burns and Crayn 2006). Other than the spots on the thighs, few differences separate the 2 species and future taxonomic actions may result in ‘L. moorei’ being placed in synonomy with L. cyclorhyncha (the older name), or relegated to subspecific rank. Further Reading. Evolutionary relationships – Tyler and Davies (1978); Maxson et al. (1982); Burns and Crayn (2006).

Genus LITORIA

Litoria nasuta (Gray, 1842) Striped Rocket Frog

!

! !

! !! ! ! ! ! ! ! ! ! !! !! ! ! ! ! !! !!! ! ! !! !!!! ! !! !!!! ! !! ! ! !

!!

Kununurra (P. Doughty)

Description. This is a moderate-sized species but remarkable in that it is so exceptionally elongated. The body is very slender and the head an elongated triangle with a prominent protruding snout. The hindlegs are extremely long (TL/S-V 0.64–0.82) and the fingers and toes long and slender. The fingers are unwebbed and bear very small terminal discs. The webbing between the toes is reduced, not reaching the ends of any of the toes. The skin of the back is raised into longitudinal folds. The background colour of the back is a mixture of shades of brown. There is a very dark brown stripe along the side of the head from the nostril to the insertion of the forearm. Males 33–45 mm; females 36–55 mm S-V. Distribution. Kimberley region. South to Beagle Bay, Mount Bell, Mount Barnett Station and Kununurra. Extends south to Mandora along 80-mile Beach. Outside of Western Australia it extends eastwards and southwards in a continuous arc to New South Wales. Also occurs in southeastern New Guinea. Habitat. Grasslands and also a variety of terrestrial habitats, dispersing widely during the

wet season. Advertisement Call. Complex: usually commences with one or two short notes ‘but ... but’, followed by a long series of ‘wick…wick…wick’ (resembling a duck or PacMan), with the short notes given intermittently between the main call. Breeding Biology. Breeds in static water. Eggs are laid in small clumps of 20–100. The tadpole reaches a length of up to 61 mm, and the teeth are long and in 2 upper and 3 lower rows; the innermost rows are narrowly divided in the middle. The tail is long with a moderately arched fin. Quite early in development the head elongates to the shape characteristic of the frog. Etymology. nasutus means ‘large-nosed’, alluding to the elongate snout. Remarks. A remarkably proficient jumper owing to its long legs and an evolutionary extension of the intercalary cartilage in the toes (Tyler 1994). Further Reading. Breeding biology – Anstis (2002).

79

FROGS OF WESTERN AUSTRALIA

Litoria pallida Davies, Martin and Watson, 1983 Pale Rocket Frog

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Kununurra (B. Maryan)

Description. A small to medium-sized, elongatebodied ground-dwelling frog. The head is angled and elongate. The fingers are long and pointed with webbing at the base. The hindlimbs are long (TL/S-V 0.56–0.68) and the toes incompletely webbed. The skin on the back is smooth or only slightly bumpy. There is a distinct stripe on the side of the head, yellow on the upper lip and the markings on the thighs are distinct and suffused with yellow in life. Background colour grey or brown. Males 24–38 mm; females 30–41 mm S-V. Distribution. Kimberley region. South to Bohemia Downs and Billiluna. Widely distributed across northern Australia in the Northern Territory and Queensland. Habitat. Temporarily flooded grasslands.

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Advertisement Call. Complex: a long highpitched wavering series of notes, with short calls given intermittently. Males call from open areas within 1 m of water. Breeding Biology. Eggs are laid on the surface of water of temporary pools in clutches of 96– 330. Ovum diameter is 1.3–1.4 mm and capsule diameter 5.8–6.8 mm (Tyler et al. 1983a as ‘Litoria sp. nr latopalmata’). Etymology. pallida refers to the ‘pale’ appearance. Remarks. Similar to the closely-related L. inermis, but L. pallida is larger, has a more pointed snout and the main note always has a waver. Further Reading. Call analysis – Tyler et al. (1983b); breeding biology – Tyler et al. (1983a).

Genus LITORIA

Litoria rothii (De Vis, 1884) Northern Laughing Tree Frog

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Mornington Station (P. Doughty)

Description. A moderate-sized species with a flattened appearance and protruding eyes. The limbs are moderately long (TL/S-V 0.48–0.53). The fingers have broad lateral fringes, distinct webbing and bear expanded terminal discs. The toes are fully webbed. Individuals are capable of a striking colour change from rich, mottled brown to a uniform and very pale cream or almost white. A distinctive feature is the highly contrasting black and yellow markings in the groin and on the posterior surface of the thigh. The supratympanic fold is black-edged. The upper half of the iris is rich red and the lower half a pale metallic gold. Males 37–50 mm; females 45–49 mm S-V. Distribution. Kimberley region. South to Coulomb Point, Dampier Downs, Napier Range, Lake Gladstone and Lissadell Station. Also across northern Australia and the southern lowlands of New Guinea. Habitat. Occurs near water, especially swamps where it will shelter in Pandanus trees. Also a common inhabitant of human dwellings.

Advertisement Call. A loud descending trill that resembles rather raucous laughter. Males call from trees or low shrubs, up to 2 m or more above the ground or from the water’s edge. Breeding Biology. Spawns at the commencement of the wet season in October-November and tadpoles can be found as late as February in temporarily flooded areas in low-lying country. Spawn is deposited in small clumps. Ovum diameter is 1.3–1.4 mm and capsule diameter 4.6–5.5 mm. In early stages tadpoles are pale yellow on top. Tadpoles become mottled brown and very large (> 10 cm). Etymology. Named after Henry Ling Roth (1855–1925), a well-known ethnologist who collected the specimens used in the description. It is also possible that his brother, Walter Roth, a doctor who worked on ethnology in north Queensland, was also involved in the collection of the type series. Further Reading. Call, reproduction and tadpoles – Martin et al. (1979).

81

FROGS OF WESTERN AUSTRALIA

Litoria rubella (Gray, 1842) Little Red Tree Frog

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Kununurra (J. Francis) Mount Brockman (P. Doughty)

surviving in arid areas by avoiding exposure to drying conditions, principally by sheltering in crevices in large rocks or other well-insulated places. Very common in and around human dwellings. Advertisement Call. A long harsh vibrant and very loud screech. A chorus is similar to a group of sea gulls. Breeding Biology. Small circular clumps of freefloating spawn up to 4 cm diameter including Description. A small and rather robust species 40–300 eggs are laid on the surface of static water with a small short head and legs (TL/S-V after summer rains. Ova are approximately 1 mm 0.34–0.43). The fingers are short and bear small diameter and the capsules 2 mm. Larval life is brief but distinct dilated discs with webbing only with metamorphosis within 40 days and possibly at the base. The toes are about three-quarters occupying only 14 days in warmer regions. webbed. The dorsal surface is reddish-brown, Tadpoles attain a maximum length of 50 mm. ranging from a very pale shade, rich red or an Etymology. rubella means ‘red’ in reference to extremely dark brown. There is usually a broad the typical colouration. dark stripe along the side of the head and body. The back is occasionally flecked with gold or black. Remarks. Litoria rubella avoid excessive water loss by resting in a tightly huddled posture, and Males 28–37 mm; females 33–43 mm S-V. by a dramatic colour change from the typical Distribution. Kimberley region and arid zone. gray, brown or fawn, to a bright white. This In the Kimberley south to Anna Plains, Edgar allows them to bask in direct sunlight in habitat Range, Lake Gladstone and Denison Range. In that provides little shelter. Owing to its common the arid zone, centred in the Pilbara region from occurrence in toilet blocks, it is sometimes Port Hedland southwest to Shark Bay and inland nicknamed the ‘dunny frog’. to Goldsworthy, Rudall River, Lake Carnegie and Cue. Elsewhere occupies the northern twoFurther Reading. Call description – Lindgren and thirds of the continent. Main (1961); breeding biology – Tyler et al. (1983a), Habitat. This ubiquitous species is capable of Anstis (2002); physiology – Withers (1995b). 82

Genus LITORIA

Litoria splendida Tyler, Davies and Martin, 1977 Magnificent or Splendid Tree Frog

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Kununurra (J. Francis)

Description. A very large tree frog. There is a large and protuberant skin gland upon the entire upper surface of the head. The fingers are broadly fringed with extremely large expanded discs and the toes are extensively webbed. The limbs are moderately long (TL/S-V 0.46–0.50). The colour is olive or green, sparsely and dramatically spotted with sulphur markings. The sulphur markings may be circular and edged with black. The hidden portions of the limbs and flanks range from bright yellow to brilliant orange. Males to 106 mm; females to 118 mm S-V. Distribution. Kimberley region. South to Kimbolton, Napier Downs and Purnululu National Park. Also extends into the Northern Territory. Habitat. Found on rocks and trees and in caves on massive sandstone rock outcrops. Advertisement Call. A slow deep bark or buzz that rises slightly in pitch.

Breeding Biology. Males call from rocks several meters from water after heavy rains in the wet season. In captivity, up to 6400 eggs have been laid by an individual. Etymology. splendida refers to the appearance of this frog. Remarks. This charismatic species is the largest Western Australian frog. It is usually very docile when handled, usually looking up for somewhere to jump. The secretions of the vast gland upon the head have been found to have antibiotic and analgesic peptides present (Bowie et al. 1998). Some of the peptides have been found in the tadpoles of this species (Wabnitz et al. 1998). Further Reading. Call analysis – Watson and Gerhardt (1997).

83

FROGS OF WESTERN AUSTRALIA

Litoria staccato Doughty and Anstis, 2007 Chattering Rock Frog

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♂ The Grotto (P. Doughty) ♁ The Grotto (P. Doughty)

Description. A small to medium-sized slender elongate frog. The head is angled and elongate with protruding eyes. The fingers have wide terminal discs and the hindlimbs are moderately long (TL/S-V 0.46–0.55) with nearly complete webbing between the toes (no webbing on fifth toe tip) and moderately expanded discs. The background colour ranges from a dark red to grey to pale beige. There is a diffuse loreal stripe that continues along the sides. A paler snout patch extends between the eyes and there are diffuse longitudinal stripes along the back. Males 26–33 mm; females 33–39 mm S-V. Distribution. Kimberley region. From the

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Prince Regent River Nature Reserve north to the Mitchell Plateau and east to near Kununurra. Also occurs on many islands. Habitat. Found on sandstone escarpments and platforms and on rocky ridges far from flowing water. Advertisement Call. Complex: the main call is a high-pitched Morse code-like call with occasional trills, interspersed with softer ‘chirps’ between the main calls. Breeding Biology. Males call near flowing streams or near seepages on steep slopes. Tadpoles are broad-bodied and show adaptations to flowing streams. They are finely stippled, reach a maximum length of 52 mm and metamorphose in about 7 weeks. Etymology. staccato is in reference to the halting Morse code-like call of this species. Remarks. This species differs only subtly in morphology from L. coplandi (i.e. smaller size, diffuse loreal and dorsal stripes and reduced webbing), but the calls are very different.

Genus LITORIA

Litoria tornieri (Nieden, 1923) Black-shinned Rocket Frog

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Mitchell Plateau (M. Anstis)

Description. A small to medium-sized slender elongate ground-dwelling frog. The head is angled and elongate, the fingers are long and unwebbed and hindlimbs long (TL/S-V 0.49– 0.67) with reduced webbing between the toes. The head and back range from a pale sandy brown to a rich blood red. There is a clearly demarcated dark brown stripe on the head that ends past the tympanum, with usually smaller spots past the arm on the sides. There is banding of brown and yellow along the length of the back of the thigh and a narrow dark edge to the tibia. Males 29–34 mm; females 33–38 mm S-V. Distribution. Kimberley region. South to Yampi Peninsula, Napier Downs and Gibb River Station. It also occurs in the northern part of the Northern Territory. Habitat. Found near the edge of shallow permanent swamps and flooded areas. Advertisement Call. Complex: usually a short

series of high-pitched notes repeated frequently, with intermittent softer calls between the main calls. In large choruses, the high-pitched notes are repeated constantly. Breeding Biology. Males call from cover within 3 m of water. Eggs are laid in large clumps in temporarily flooded areas. The ovum diameter is 1.4 mm; the egg capsule diameter is 3.2–3.7 mm. The tadpoles have long, pointed tails and complete their development in 6–7 weeks. Etymology. Named after Gustav Tornier (1858– 1938), a colleague of Fritz Nieden at the Berlin Museum and who is best known for his work on African frogs. Tornier retired as the Director of the Museum the year this species was described. Remarks. Similar to the other ground frogs L. inermis and L. pallida, from which it can be distinguished by its reddish colour and the black stripe on the shin.

85

FROGS OF WESTERN AUSTRALIA

Litoria watjulumensis (Copland, 1957) Wotjulum Frog

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Mornington Station (J. Francis)

Description. A medium to large-sized elongate frog of variable appearance and very long limbs (TL/S-V 0.59–0.73). The fingers are long, slender, unwebbed and bear distinct terminal discs. The toes are fully webbed and have enlarged, circular, terminal discs. Most commonly individuals are a uniform pale brown dorsally, but they may be very densely mottled with intense dark brown or with grey. There is a broad and clearly demarcated dark brown stripe passing from the nostril to the eye; broadening behind the eye and extending to above the insertion of the forearm. The flanks are often suffused with vivid pale yellow. Males 33–41 mm; females 42–70 mm S-V. Distribution. Kimberley region. South to Yampi Peninsula, Noonkanbah and Purnululu National Park. Also in the northern portion of the Northern Territory and northwest Queensland. Habitat. Most commonly found at the edges of permanent or semi-permanent slow-flowing rocky creeks, but also occurs in flooded grasslands during the wet season. Advertisement Call. Very complex. Sporadic chicken-like ‘clucks’ are given, followed by a rapid series of evenly-spaced notes (3–4 per second) which adruptly double in rate without a pause for up to 30 seconds, and is again followed

86

by more ‘clucks’. The ‘clucks’ are usually given in response to other males’ calls, and the series of rapid notes are usually given in unison. Males call from open areas at the edge of static water or slow-flowing rocky creeks, only earnestly beginning to call well after midnight and continuing into the first hours of daylight. Breeding Biology. Floating clumps of 30–200 eggs are laid in temporary pools. Ovum diameter range is 1.6–1.9 mm and the capsules 4.2–4.9 mm. The tadpole is mottled with dark brown and attains a maximum length of 42 mm. Larval life occupies approximately 7–8 weeks. Etymology. The name of this species is derived from the now abandoned Wotjulum Mission on Yampi Sound. Unfortunately, Copland misspelled Wotjulum in his original description (he used an ‘a’, not an ‘o’) which is not correctable owing to the rules of naming which are focused on stability. Remarks. Litoria watjulumensis is remarkable not only for its call but pronounced sexual size dimophism with some females twice as long as males and many times more massive. The expanded discs on the digits and preference for rocky creeks may indicate a closer relationship to the rock frogs (L. coplandi and L. staccato) than the other ground frogs (L. inermis, L. pallida, L. nasuta, L. tornieri).

Genus LITORIA Litoria inermis

Litoria watjulumensis

Litoria pallida

Litoria tomieri

Litoria nasuta

Sonograms of ground frogs of the genus Litoria. The time interval for all calls (except for L. watjulumensis) is 4 seconds along the X-axis; the Y-axis represents amplitude. The call of L. watjulumensis is more sustained and the X-axis is 20 seconds. Calls of all species can be heard on the Alcoa Frog Watch web site – http://frogwatch.museum.wa.gov.au). 87

Genus METACRINIA Parker, 1940 Diversity and Evolutionary Relationships A monotypic genus of myobatrachine frogs found only in southwestern Australia. Genetic evidence suggests a sister relationship with Arenophryne and Myobatrachus (Roberts et al. 1997; Read et al. 2001). Metacrinia lacks the overt morphological specializations of the two sand-adapted genera, except that they all share forwards crawling/burrowing and have large, direct-developing eggs.

Morphological Characteristics Head small; body depressed; tympanum present; jaws lack teeth; tongue slender and elongated. Limbs very short; fingers unwebbed; first finger shorter than second. Crawls rather than hops.

Reproductive Mode Terrestrial reproduction. Eggs large and laid out of water; subsequent process of larval development occurs entirely within the egg capsule with a miniature of the adult emerging several months later (Anstis 2008).

Etymology Meta means ‘near’ in reference to a perceived superficial similarity to Crinia.

Genus METACRINIA

Metacrinia nichollsi (Harrison, 1927) Forest Toadlet

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Dunsborough (G. Harold)

Description. A small short-bodied species with short hindlegs (TL/S-V 0.24–0.34) and short fingers and toes. Tympanum visible. This species is brilliantly marked with vivid orange patches in the groin, behind the thighs and upon the upper arm contrasting with the purplish or dark slate background colour. Undersurface heavily pigmented with slate or dark purple, often bearing yellow or orange marks at the insertion of the limbs. Males 19–21 mm; females 22–28 mm S-V.

Distribution. Southwest region. Restricted to the karri belt in the extreme southwest from Dunsborough south and east to Albany, with a population in the Stirling Ranges. Habitat. Found in leaf litter and under logs in karri forests. Advertisement Call. A short grating squelch. Breeding Biology. A direct developer with the larval period (> 2 months) occurring entirely within the egg capsule. Egg capsule diameter is approximately 5 mm. Approximately 12–30 eggs under cover such as beneath a log in late summer. Etymology. Named after G. E. Nicholls of the University of Western Australia (1921–1947), an early contributor to amphibian systematics. Further Reading. General – Barbour and Loveridge (1929), Main (1965a), Driscoll (1998c); reproduction – Harrison (1927), Anstis (2008).

Metacrinia nichollsi, Walpole (B. Maryan) 89

Genus MYOBATRACHUS Schlegel, 1850 Diversity and Evolutionary Relationships A highly distinctive genus of myobatrachine frogs restricted to southwestern Australia. The closest relative is Arenophryne, with which it shares a number of osteological features, the habit of head-first burrowing with the hands, and a reproductive system based on direct development.

Morphological Characteristics Head small, body bulbous, eyes very small and inconspicuous; tympanum not readily detectable. Limbs very short; fingers short, broad and unwebbed; toes unwebbed.

Reproductive Mode Direct development, i.e. all embryonic stages are passed entirely within the egg. Eggs very large, unpigmented.

Etymology Myo means ‘muscle’ and batrachian refers to frogs, hence ‘muscle-frog’ probably in reference to the powerful limbs.

Genus MYOBATRACHUS

Myobatrachus gouldii (Gray, 1841) Turtle Frog

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Kellerberrin (B. Maryan)

Description. A rather flattened small to moderately large frog with a very small head, tiny eyes, very short but muscular limbs (TL/S-V 0.20–0.24) and reduced but very broad fingers and toes. The limbs tend to extend from the sides of the body rather than beneath it. This feature and the slight hump in the middle of the back produces the turtle-like appearance that leads to the common name of the species. The back varies from yellow-pink to dull slate to dark brown. Males 26–42 mm; females 27–51 mm S-V. Distribution. Southwest region and adjacent arid zone. From Kalbarri National Park in the north, south and east to Morowa, Quairading, Southern Cross and Grass Patch. Absent from the jarrah-karri belt. Habitat. Occurs most commonly in sandy soils. Advertisement Call. The advertisement call is a series of deep croaks made on the surface but males may call from under the surface early in the breeding season.

Breeding Biology. Males call from September to February. Males and females pair up and then dig a deep burrow and remain together until autumn – one of the rare instances of extended pairing in a frog species. Clutches of 9–38 eggs are laid 0.5–1.5 m underground. The eggs are very large (capsules to 7.4 mm). Hatchlings range from 9–11 mm and are miniatures of the adults, including the ability to burrow. Etymology. Named after John Gould (1804– 1881), famous ornithologist and artist who sent specimens to the British Museum that were probably collected by John Gilbert. Remarks. A specialist feeder on termites. Further Reading. General – Philipp (1958); evolutionary relationships – Tyler et al. (1980), Davies (1984), Maxson and Roberts (1985), Read et al. (2001), Frost et al. (2006); diet – Calaby (1956); reproduction – Watson and Saunders (1959), Roberts (1981), Anstis et al. (2007); physiology – Withers and Roberts (1993).

Myobatrachus gouldii, Three Springs (G. Harold) 91

Genus NEOBATRACHUS Peters, 1863 Diversity and Evolutionary Relationships A complex genus of limnodynastine frogs, with 10 species distributed across most of southern and inland Australia, extending into the arid northwest of Western Australia. Eight species occur in Western Australia, with up to 4 or 5 within a region, except the Kimberley. Evolutionary relationships within the Limnodynastidae are still unresolved, with some support for a sister relationship between Neobatrachus and Heleioporus (Roberts and Watson 1993) or with Neobatrachus as the sister taxon to Notaden, Platyplectrum and Lechriodus (Frost et al. 2006). Neobatrachus is unique among Australian frogs in the occurrence of polyploidy (duplicate copies of chromosomes). Four of the 10 species are tetraploid, that is they have 4 sets of chromosomes instead of 2 (usually 1 inherited from each parent). Polyploidy in vertebrates occurs most commonly as a consequence of infrequent hybridization. The tetraploid species are N. aquilonius, N. centralis, N. kunapalari and N. sudelli, the last of which is restricted to eastern Australia but may prove to be a senior synonym of N. centralis. Evolutionary relationships among diploid and tetraploid species were examined biochemically by Mable and Roberts (1997) and with call structure analysis by Roberts (1997a,b). Their results converge in suggesting a close relationship between the tetraploids N. centralis and N. sudelli, which may be better treated as clinal geographic variants of the same species. Here we maintain a conservative stance and treat them as valid species until further evidence becomes available.

Morphological Characteristics Head and body robust; pupil vertical (Figure 2A); tympanum visible; upper jaw toothed; tongue broad. The skin is often loose-fitting, especially near the hindquarters. Limbs short; fingers unwebbed; first finger longer than second. Toes partly webbed; inner metatarsal tubercle large, shovel-shaped and either black or unpigmented.

Reproductive Mode Aquatic reproduction. Eggs pigmented; laid in long strings, often attached to emergent or submerged aquatic vegetation. Larval life in several species occupies about 6–7 weeks.

Etymology Neo means ‘new’ and batrachos refers to frogs, hence ‘new frog’.

Remarks Most, and possibly all, species of Neobatrachus can form a protective skin ‘cocoon’ as a means of reducing water loss during periods of aestivation.

Further Reading Cocoon formation and physiology – Lee and Mercer (1967), Withers (1993, 1995a, 1998).

Neobatrachus kunapalari, Leinster (G. Harold) 93

FROGS OF WESTERN AUSTRALIA

Neobatrachus albipes Roberts, Mahony, Kendrick and Majors, 1991 White-footed Trilling Frog

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Ravensthorpe (G. Harold)

Description. A medium-sized Neobatrachus with short limbs (TL/S-V 0.30–0.37). Toes with moderately extensive webbing (Figure 23). Inner metatarsal tubercle pale brown or white; no outer metatarsal tubercle. Back brown with poorlydefined darker markings sometimes including an incomplete vertebral stripe. There is a broad pale ‘V’, sometimes reduced to a spot, between the eyes. Ventral surface of body, legs and upper arms creamy-white. Skin of upper surface of foot is white. Males 33–46 mm; females 35–43 mm S-V. Distribution. Eastern southwest region and the southwest arid zone. Eastern wheatbelt and adjacent goldfields from Wyalkatchem, Bruce Rock, Narembeen, Quairading, Dongalocking and the Stirling Ranges in the west, east to Coolgardie and Cape Arid and south of the Great Eastern Highway. Also at Junanan Rock. Habitat. Temporary water bodies in agricultural regions. Advertisement Call. A trill of 36–40, short, rapidly repeated pulses. Breeding Biology. Most activity has been

94

recorded after autumn rains but males have also been heard after rain in October, March and January. Males call from covered sites and deep water (30–50 cm deep or more). Etymology. albipes means ‘white-footed’. Remarks. This species is most likely to be confused with N. pelobatoides which also has reduced pigment on the inner metatarsal tubercle and a tendency to have a vertebral stripe (more prominent and consistent in N. pelobatoides).

Figure 23

Sole of right foot of Neobatrachus albipes showing extent of webbing and form of inner metatarsal tubercle.

Genus NEOBATRACHUS

Neobatrachus aquilonius Tyler, Davies and Martin, 1981 Northern Burrowing Frog

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Warree Hill (B. Maryan)

Description. A medium-large robust-bodied frog which is generally bright yellow or greenish, bearing extensive, darker (usually blackish) irregular-shaped markings. The limbs are very short (T/S-V 0.29–0.35) and their undersurfaces are plum. Toes with moderately extensive webbing. Ventral surface of body white. Males 48–54 mm; females 52–59 mm S-V. Distribution. Arid zone and southwestern edge of Kimberley region near Broome and Derby. Recorded from scattered localities in the arid zone, from the coastal Pilbara, western Gibson Desert and south to Kirkalocka, Yamana and Gidgie. Also occurs in the Northern Territory. Habitat. Found in sparsely vegetated country. Active at the edge of the water. Advertisement Call. A short, low trill repeated at frequent intervals.

Breeding Biology. An opportunistic and ‘explosive’ breeder emerging to breed in response to cyclonic rains. Eggs are laid in shallow flooded areas. Females may lay up to 1500 eggs. Etymology. aquilonius means ‘northern’ (technically ‘north-eastern’) in reference to this species being the most northerly occurring member of the genus Neobatrachus. Remarks. This species is a tetraploid (i.e. has double the number of chromosomes). Further Reading. Call analysis – Tyler et al. (1981b), Roberts (1997a,b); physiology and burrowing – Withers (1995a, 1998), Thompson et al. (2005), Cartledge et al. (2006b).

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FROGS OF WESTERN AUSTRALIA

Neobatrachus centralis (Parker, 1940) Desert Trilling Frog

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Leinster (G. Harold)

Description. A moderate-sized robust-bodied Neobatrachus with short limbs (TL/S-V 0.28– 0.37). Breeding males have small black spines on the back. They are generally dull bronze, gold or yellow, with irregular dark brown blotches. Males 41–49 mm S-V. Distribution. Arid zone. In Western Australia known only from near Mount Magnet with confidence where calling males were karyotyped. Their ill-defined range extends east through the southern arid zone. Habitat. Unknown. Advertisment Call. A short high-pitched trill, distinguishable from N. kunapalari by lower pulse number and higher frequency. Breeding Biology. Tadpoles in the eastern states are up to 70 mm with a wide body and shallow fins. They take about 40 days to complete development. Etymology. centralis refers to this species’ central

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distribution on the Australian continent. Remarks. This species was originally described from a site near Lake Eyre, South Australia and was, until recently, considered to be common and widespread in Western Australia. Now that it is no longer confused with other Neobatrachus species in Western Australia, its distribution appears to be far less extensive. Further work is necessary, but is hampered by the distance and lack of sealed roads making access to breeding ponds near the type locality difficult when it rains in the arid zone. Tetraploid chromosome number. This species may be a junior synonym of N. sudelli Lamb, 1911 (Roberts 1997a). Further Reading. Type locality – Tyler and Ledo (1973); call analysis – Roberts (1997a,b); diet and demography – Read (1999a,b); tadpoles – Main (1968), Davies (1991), Anstis (2002); physiology and cocoon formation – Withers (1993, 1995a, 1998), Fuery et al. (1998).

Genus NEOBATRACHUS

Neobatrachus fulvus Mahony and Roberts, 1986 Tawny Trilling Frog ! ! ! ! !!! ! ! !

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Wooramel (G. Gaikhorst)

Description. A medium-sized robust Neobatrachus with short limbs (TL/S-V 0.28– 0.40). Skin on side of body extends across to knee so that the groin is indistinct. Toes with moderately extensive webbing. Breeding males without spines on back. They have a chocolate back interrupted by yellow spots and variegations. Males 38–42 mm; females 42–48 mm S-V. Distribution. Arid zone. A narrow band on the west coast from Shark Bay north to the North West Cape. Habitat. Found on claypans, red soils and

sandy brown loams with mulga, soft grasses and Triodia. Advertisement Call. A short trill similar to N. aquilonius. Breeding Biology. Calls and breeds after cyclonic rains in summer and in early winter. Etymology. fulvus means ‘reddish yellow’ or ‘tawny’. Further Reading. Call analysis – Roberts and Majors (1993); physiology – Withers (1993, 1995a, 1998).

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FROGS OF WESTERN AUSTRALIA

Neobatrachus kunapalari Mahony and Roberts, 1986 Wheatbelt Frog

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Ponier Rocks (B. Maryan)

Description. A medium-large robust-bodied Neobatrachus with short limbs (TL/S-V 0.32– 0.36). Skin extending only slightly from side of body to encompass groin. Toes with moderately extensive webbing. Breeding males have small black spines on the back. The background colour is a pale yellow to tan with many irregular pale and dark brown patches on the back. Males 48–58 mm; females 53–61 mm S-V. Distribution. Southwest region and arid zone south of Menzies and Wubin. Extends to the edge of the Nullarbor Plain. Habitat. Usually found on impervious soils where pools often contain a milky, opaque colloidal mixture, effectively hiding tadpoles from view.

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Advertisement Call. A long, low trill. Breeding Biology. An opportunistic breeder emerging and spawning during heavy rainfall at any time of the year. Several males may attempt to mate with a female when she enters a breeding chorus. Etymology. kunapalari means ‘frog’ in the language of the Gugadja people. Remarks. A tetraploid species with double the normal number of chromosomes. Further Reading. Call analysis – Littlejohn and Main (1959; as N. centralis), Roberts (1997a,b); breeding behaviour – Davis and Roberts (2004); tadpoles – Davies (1991); physiology – Withers (1993, 1995a, 1998).

Genus NEOBATRACHUS

Neobatrachus pelobatoides (Werner, 1914) Humming Frog

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Ellenbrook (B. Maryan)

Description. A medium-sized species with a slightly rotund appearance, moderately short limbs (TL/S-V 0.34–0.41) and protruding eyes. The dorsal skin is more finely tubercular than in the other members of the genus and the toes have less extensive webbing (Figure 24). The colour is generally dull green or pale yellow, extensively covered with clearly defined, irregularly shaped patches of dark brown or green. There is usually a narrow red mid-dorsal stripe along the middle of the head and body. Males 37–45 mm; females 36–44 mm S-V. Distribution. Southwest region and adjacent arid zone. From Nerren Nerren, Perenjori, Dalwallinu, Mount Elvire and Ponier Rocks in the north and east; south to Buckingham, Stirling Ranges and Israelite Bay. Habitat. Generally associated with clay or loam soils; occurs near granite outcrops and farm dams in the wheatbelt. Advertisement Call. A long low trill or ‘hum’, heard only over a short distance. Breeding Biology. Breeding takes place from

May to July in rock pools, ditches and farm dams. Etymology. pelobatoides means ‘resembling Pelobates’, a genus of European frogs. Further Reading. Call analysis – Littlejohn and Main (1959), Main (1965a), Roberts (1997b); tadpoles – Main (1965a); physiology – Withers (1993, 1995a, 1998).

Figure 24

Sole of right foot of Neobatrachus pelobatoides showing less extensive webbing of toes than other species of Neobatrachus (compare with Figures 23 and 25).

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FROGS OF WESTERN AUSTRALIA

Neobatrachus sutor Main, 1957 Shoemaker Frog !

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Twin Peaks Station (B. Maryan) Central Ranges (P. Doughty)

Description. A moderate-sized rotund very shortlegged (TL/S-V 0.27–0.37) frog with prominent eyes, a high rounded snout and extensively webbed toes. The inner metatarsal tubercle is heavily pigmented and sharply keeled. The back is brilliant gold to dull red, bearing numerous small and/or irregular patches of rich brown or black. Males 35–45 mm; females 34–51 mm S-V. Distribution. Arid zone and adjacent southwest region. Upper west coast from North West Cape south to Wooramel River and the mid-west interior from Jiggalong in the north, south to Lake King. West to Mingenew and Mullewa, Fraser Range, Queen Victoria Spring and Warburton Range. An apparently isolated population in the central wheatbelt near Dongolocking. Also occurs in adjacent areas of the Northern Territory and South Australia. 100

Habitat. Areas of clay, loam or sandy soils. Advertisement Call. A repetitive tapping resembling the sound of a shoemaker’s hammer at work. Breeding Biology. A summer breeder (February to March) that responds to cyclonic rains. Eggs are laid in static water in claypans. Development is rapid, with metamorphosis occurring around 40 days. Etymology. sutor means ‘cobbler’ in reference to the sound of the call which resembles a shoemaker’s tapping. Further Reading. Physiology – Withers (1993, 1995a, 1998)

Figure 25

Sole of right foot of Neobatrachus sutor showing extent of webbing and form of inner metatarsal tubercle.

Genus NEOBATRACHUS

Neobatrachus wilsmorei (Parker, 1940) Plonking Frog

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Perenjori (B. Maryan)

Description. A medium-large robust frog with very short limbs (TL/S-V 0.30–0.34) and moderately extensive webbing between the toes. The back is dark brown and bears 5 narrow and straight, bright yellow stripes along its body forming a large ‘W’. There is a line extending from the tip of the snout to a position behind the thighs, and on each side of the body a pair of diverging lines commencing behind the head, so forming a W-shaped mark. One of these lines passes slightly upwards towards the coccyx, while the other runs parallel to the midline and terminates in front of the groin. Males 56–61 mm; females 51–63 mm S-V. Distribution. Arid zone. On the west coast from the Lyndon River south to the Irwin River and east to Lake Nabberu, Banjawarn and Yundamindra in the north; south to Carnarvon, Gnoolowa Hill, Morowa, Paynes Find and Kalgoorlie.

Habitat. Inhabits areas subjected to seasonal flooding. Advertisement Call. A hollow ‘plonk’ repeated slowly. Breeding Biology. Breeds after summer rains. The tadpoles complete development in about 40 days. Etymology. Parker did not indicate who he named species after in his 1940 review. This species may have been named after Norman T. Wilsmore (1868–1940), a chemist at the University of Western Australia who was once based in London. Further Reading. Call analysis – Littlejohn and Main (1959); tadpoles – Davies (1991); physiology – Flanigan et al. (1993), Withers (1993, 1995a, 1998).

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Genus NOTADEN Günther, 1873 Diversity and Evolutionary Relationships A small but distinctive genus of ball-shaped limnodynastine frogs. The 4 species’ combined range extends across the tropical north and through the arid zone into inland eastern Australia. One species is endemic to the Kimberley, 1 centred in the arid zone and 1 in tropical Australia. All 3 are found in different parts of the Kimberley, but never together. Phylogenetic affinities may be with Heleioporus and Neobatrachus (Roberts and Watson 1993) or Platyplectrum and Lechriodus (Frost et al. 2006).

Morphological Characteristics Head rather small; body large and globular; pupil horizontal with ventral extension; tympanum not visible; upper jaw without teeth; tongue broad, but elongated when extended. Dorsum and flanks with numerous small, rounded warts, sometimes coalescing to form longitudinal ridges. When aroused, all species exude a viscous, white or cream secretion from their glandular skin. Limbs very short; fingers unwebbed (Figure 11A); first finger longer than second. Toes unwebbed; inner metatarsal tubercle large and shovel-shaped, black or unpigmented.

Reproductive Mode Aquatic reproduction. Eggs pigmented; spawn laid in large clumps in shallow water. Tadpoles are freeswimming. Larval life is probably short with metamorphs of one species measuring only 13–14 mm.

Etymology From notos meaning ‘back’ and aden meaning ‘gland’.

Remarks All species give a loud ‘whoop’ as an advertisement call and all can exude a milky viscous substance when handled. This secretion is being investigated as a possible surgical adhesive (Graham et al. 2005, 2006).

Notaden melanoscaphus, Kununurra (P. Doughty) 103

FROGS OF WESTERN AUSTRALIA

Notaden melanoscaphus Hosmer, 1962 Northern Spadefoot

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Mornington Station (J. Francis)

Description. A robust medium-sized species with a circular body and very short legs (TL/S-V 0.29–0.32). The skin is of an exceptionally rough texture. The hands and feet bear long pointed digits, and there is a sharp black shovel-shaped inner metatarsal tubercle. Dorsal colour ranges from an intense and predominantly uniform slate, to pale grey bearing a dark brown patch linking the upper eyelids and irregular dark brown patches in a double row down the back. Males 34–54 mm; females 45–49 mm S-V. Distribution. Kimberley region. From Truscott south to Mornington Station and east to Lake Argyle. Widespread across northern Australia in the Northern Territory and Queensland. Habitat. A burrowing species associated with impervious clay soils. Advertisement Call. A loud ‘whoop’, repeated frequently.

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Breeding Biology. Breeds in shallow flooded areas of sparse, low grasses upon clay soils. Males call while floating in the water with their lungs and vocal sacs distending their volume about threefold. Spawn is laid in a large submerged mass of up to 1200 eggs. The tadpole is most unusual with a small mouth lacking upper labial teeth; they feed on suspended particles. Etymology. melano means ‘black’ and scaphus means ‘spade’ in reference to the black metatarsal tubercle diagnostic of this species. Remarks. When handled, specimens release from their glandular skin a copious secretion of pale cream exudate which rapidly turns bright orange and hardens upon collectors’ hands. It resembles a contact adhesive in its consistency, and requires the use of abrasive soaps for removal.

Genus NOTADEN

Notaden nichollsi Parker, 1940 Desert Spadefoot !!! ! ! !! ! !

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De Grey River (B. Maryan)

Description. A medium to large, round-bodied species with very short legs (TL/S-V 0.28–0.33). The fingers and toes are long and pointed, and the inner metatarsal tubercle is shovel-shaped and unpigmented. Most individuals are a dull orange, olive-grey or brown upon which there are groups of raised black tubercles (sometimes tipped with red or yellow) in a longitudinal pattern. Males 42–63 mm; females 46–67 mm S-V. Distribution. Arid zone and southwest Kimberley region. From Derby east to vicinity of Billiluna and south to Karratha, Mount Edgar, Jiggalong and Alexandra Spring. Also at Yalabia near Carnarvon and Mount Linden. Also occurs in the Northern Territory. Habitat. Open country with sparse vegetation. Prefers to burrow in sand dunes. Advertisement Call. The call is a loud ‘whoop’, repeated frequently. Males usually call while floating in the water.

Breeding Biology. An opportunistic breeder. Eggs are laid in temporarily flooded areas of clay or sandy soils. The spawn is a large, shapeless clump of over 1000 eggs. Tadpoles have 3 upper and 3 lower tooth rows. Larval life may be completed within 30 days and newly metamorphed frogs are 13–14 mm in length. Etymology. Named after G. E. Nicholls of the University of Western Australia (1921–1947), an early contributor to amphibian systematics. Remarks. This burrowing species is active upon the surface at night even under hot dry conditions. Individuals burrow to a depth of at least 1 m. Possibly an ant and termite specialist. Further Reading. Burrowing – Slater and Main (1963), Thompson et al. (2005); diet – Calaby (1960); physiology – Main and Bentley (1964), Withers (1993), Cartledge et al. (2006b).

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FROGS OF WESTERN AUSTRALIA

Notaden weigeli Shea and Johnston, 1987 Kimberley Spadefoot

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Prince Regent River NR (P. Doughty) Defensive posture (P. Doughty)

Description. A large and relatively long-legged Notaden species (TL/S-V 0.34–0.38) with raised tubercles on the skin. The back is an orangey red with numerous white-tipped orange tubercles and indistinct grey-green patches; face grey with white tubercles; belly grayish. The inner metatarsal tubercle is large and unpigmented. Males 69–79 mm; females 70–80 mm S-V.

Distribution. Kimberley region. Known from Mitchell Plateau, Prince Regent River Nature Reserve and inland to Mount Elizabeth Station. Also on Bigge Island and Augustus Island. Habitat. Acid sands associated with sandstones and supporting tussock grasses (Triodia), eucalypts and Acacia. Advertisement Call. A loud ‘whoop’ similar to other Notaden. Breeding Biology. Probably breeds in rock pools and other available water bodies in the wet season. Etymology. Named after John Weigel, of the Australian Reptile Park in Gosford, who led the field trip on which the majority of the type series was collected.

Platyplectrum spenceri, Leinster (G. Harold) 106

Genus PLATYPLECTRUM Günther, 1863 Diversity and Evolutionary Relationships A genus of only 2 species, recently removed from the genus Limnodynastes. Recent genetic work (Frost et al. 2006) has indicated a closer affinity to Lechriodus from eastern Australia and New Guinea. Although these authors transferred P. ornatum and P. spenceri to the genus Opisthodon Steindachner, 1867, Platyplectrum is the oldest available name (and not to be confused with Platyplectron Peters, 1863 – a genus synonomized with Limnodynastes). The specific name of ‘ornatus’ is now P. ornatum.

Morphological Characteristics Both forms are small to medium-sized, round burrowing frogs that occur in the tropical and arid northern half of Australia. Head rather small; body globular; pupil horizontal with ventral extension; tympanum not visible; upper jaw without teeth; tongue broad, but elongated when extended. Limbs are short to moderately long; fingers unwebbed (Figure 11A); first finger longer than second. Toes without webbing to fully-webbed; inner metatarsal tubercle large and shovel-shaped. Dorsum and flanks with numerous small rounded warts, sometimes coalescing to form longitudinal ridges.

Reproductive Mode Aquatic reproduction. Males call while floating in water and eggs are laid in a foam nest that rapidly breaks down. Tadpoles highly carnivorous.

Etymology Platy means ‘flat’ and plectrum ‘spur’; hence ‘flat-spurred’ in reference to the spade-like metatarsal tubercles of many burrowing frogs.

FROGS OF WESTERN AUSTRALIA

Platyplectrum ornatum (Gray, 1842) Ornate Burrowing Frog

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Mornington Station (J. Francis)

Description. A medium-sized globular-bodied frog with moderately long legs (TL/S-V 0.42– 0.50). The eyes are large and protuberant. The fingers and toes are elongate, cylindrical and unwebbed (at most a trace of webbing at the base of the toes). There is a large shovel-shaped inner metatarsal tubercle, and the subarticular tubercles are prominent. The colour is pale grey, sandy or pale brown, bearing an extremely elaborate pattern of darker markings of bars and patches; some individuals have a pale yellow mid-dorsal stripe. Males 31–44 mm; females 35–50 mm S-V. Distribution. Kimberley region. South to Dampier Downs, Fitzroy Crossing and Halls Creek. Also occurs in the Northern Territory,

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Queensland, New South Wales and the Torres Strait Islands. Habitat. Areas subjected to seasonal inundation. Advertisement Call. A rather nasal ‘unk’ made as males float on the surface of the water. This species has a repertoire of other noises which it produces when disturbed during calling activity. Breeding Biology. Breeds throughout the wet season from November until March. The spawn clump of approximately 1000 eggs is a fragile foam nest, which collapses within a matter of hours to form a floating layer up to 7 cm across. The tadpoles are small and black and up to 47 mm in length. Tadpoles are often predators of other species’ tadpoles. Juvenile frogs are up to 10 mm long; metamorphosis has been recorded after 25 days. Etymology. ornatum means ornately-patterned. Remarks. Platyplectrum ornatum is closely related to P. spenceri. In Western Australia the ranges of the two species are separated by the Great Sandy Desert. Further Reading. Breeding biology – Tyler et al. (1983a), Tyler and Crook (1987), Anstis (2002).

Genus PLATYPLECTRUM

Platyplectrum spenceri (Parker, 1940) Centralian Burrowing Frog ! ! ! ! !

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Nullagine (M. Peterson)

Description. A medium-sized robust species. The limbs are of moderate length (TL/S-V 0.37–0.46). Whereas P. ornatum has no webbing or, at most, a trace at the base, P. spenceri has the toes at least one-half webbed, and commonly webbed to the tips of the toes. In individuals with extensive webbing, the web tends to be thick and fleshy. The nuptial pad is depicted in Figure 14C. Males 29–51 mm; females 33–51 mm S-V. Distribution. Arid zone. From the Pilbara south to the Gascoyne region and inland to Menzies and Minilya Station. Also extensively distributed in southern Northern Territory and the northeast of South Australia.

Habitat. Occupies areas where there are sandy soils within rocky terrain. Common in sandy creek beds, emerging at dusk to feed. Advertisement Call. A rapidly repeated, short grating note. Breeding Biology. An opportunistic species that breeds in summer in response to cyclonic rains. Etymology. Named after Walter Baldwin Spencer, biologist of the Horn expedition of 1894 and several subsequent expeditions to central Australia (Spencer 1896). Further Reading. Diet – Main and Calaby (1957); physiology – Warburg (1967), Withers (1993).

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Genus PSEUDOPHRYNE Fitzinger, 1843 Diversity and Evolutionary Relationships A moderately large genus of myobatrachine frogs, with 13 species in southern and eastern Australia. In Western Australia, there are 2 species in the southwest and 1 species in the Pilbara and Gascoyne regions. The closest relatives of Pseudophryne are 3 related genera endemic to Western Australia: Myobatrachus, Arenophryne and Metacrinia (all of which crawl forwards through leaf-litter or sand and have large directdeveloping eggs)(Read et al. 2001; Frost et al. 2006). The 2 southwestern species (P. guentheri and P. occidentalis) are closely related (Roberts and Maxson 1989).

Morphological Characteristics Head small; body rather elongated; pupil horizontal with vertical extension; tympanum absent; upper jaw without teeth; tongue slender and elongated. Limbs very short; fingers unwebbed; first finger shorter than second, toes cylindrical and unwebbed; inner metatarsal tubercle not compressed, large or small.

Reproductive Mode Aquatic reproduction. Eggs large, unpigmented or with slight pigmentation of animal pole; spawn laid out of water in the 2 southwestern species, with early, intracapsular development at site of egg deposition with subsequent tadpoles in water. Eggs of P. douglasi from arid northwest reputedly deposited among rocks in shallow water which would be unique to this genus. Larval life occupies several months.

Etymology Pseudo means ‘false’ and phryne means ‘toad’, thus ‘false toad’ in allusion to the shape and wartiness of the species, similar to the European toad familiar to Fitzinger, an Austrian herpetologist.

Pseudophryne occidentalis, Newman Rocks (G. Harold) 111

FROGS OF WESTERN AUSTRALIA

Pseudophryne douglasi Main, 1964 Gorge Toadlet ! ! ! ! !

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♂ Kotka Gorge (B. Maryan) ♁ Mt. Brockman (P. Doughty)

Description. A small and rather elongate species with short hindlegs (TL/S-V 0.31–0.39) and poorly developed metatarsal tubercles. The back is smooth or tubercular, and brilliantly marked with clearly demarcated orange markings upon a brown background. Undersurface with large areas of black variegations upon a white background or entirely slate. Males 23–28 mm; females 27–32 mm S-V. Distribution. Arid zone. In the Hamersley Range of the Pilbara from Millstream to Weeli

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Wolli Spring, Barlee Ranges, Mount Augustus and the Cape Range. Habitat. Deep gorges and canyons with permanent springs or well-vegetated pools. Advertisement Call. A short harsh note typical of other Pseudophryne. Breeding Biology. Main (1964) reported finding 89 freshly-laid eggs with a female under stones in a spring. The eggs measured 4–5 mm diameter. Etymology. Named after Athol M. Douglas who collected the first specimens in 1955.

Genus PSEUDOPHRYNE

Pseudophryne guentheri Boulenger, 1882 Crawling Toadlet

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Mundijong (B. Maryan) York (S. Donnellan)

Description. A small robust-bodied frog with short hindlegs (TL/S-V 0.27–0.30) and welldeveloped metatarsal tubercles (Figure 26). The back is extensively mottled with irregularly shaped patches of grey and/or brown. There is generally a broad dark transocular bar. Undersurface with many small to large, irregular areas of grey or black on a white background. Males 26–30 mm; females 29–33 mm S-V. Distributed. Southwest region and adjacent arid zone. From near Shark Bay extending east and inland through Mount Magnet and Booanya near Mount Ragged. .

Habitat. Commonly occurs beneath ground cover such as rocks, timber and dead leaves. Associated with granite outcrops in the wheatbelt. Advertisement Call. A short grating squelch. Males call from small tunnels in the soil. Breeding Biology. Eggs are laid following rains in autumn or early winter. They are deposited in the damp breeding tunnels, with the male often remaining with the eggs. Each egg is separate and has a tough outer capsule. The early stages of development occur within the capsule, and the tadpoles emerge at an advanced stage of development when the tunnels are flooded. Tadpoles metamorphose about 2 months later. Etymology. Named after Albert Günther (1830–1914) of the British Museum who named numerous Australian species of frogs and reptiles, and was Boulenger’s immediate predecessor. The letters ‘ue’ replace the umlaut.

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FROGS OF WESTERN AUSTRALIA

Pseudophryne occidentalis Parker, 1940 Western Toadlet

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Weowanie Rock (B. Maryan) Yellowdine (M. Peterson)

Description. A small, robust-bodied frog with short legs (TL/S-V 0.27–0.32) and distinct but relatively small and widely separated metatarsal tubercles (Figure 26). The back is deep chocolate, usually with small orange or bright yellow markings on the top of the head, the upper part of the arms and on the rump. The undersurface has large irregular areas of black upon a white background. Males 22–24 mm; females 21–35 mm S-V.

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Distribution. Southwest region and arid zone. Generally distributed to the east and north of P. guentheri. Its western limits are Shark Bay, Bruce Rock and Phillips River; its eastern limits are Tallering Barr Smith Range, White Cliffs and Balladonia Rock. Habitat. Associated with granite outcrops in the arid zone and low-lying areas subjected to flooding in the wheatbelt with clay soils. Advertisement Call. A short harsh grating rasp. Breeding Biology. Generally similar to P. guentheri except that it will also breed after summer rain. Etymology. occident means ‘west’, thus ‘from the west’. Further Reading. Taxonomy – Heyer and Liem (1976), Tyler and Davies (1980).

Genus PSEUDOPHRYNE

Figure 26

Figure 27

Sole of foot of Pseudophryne guentheri. The metatarsal tubercles are larger and more closely approximated than in the otherwise similar P. occidentalis.

Sole of foot of Pseudophryne occidentalis showing size and degree of separation of metatarsal tubercles.

Pseudophryne guentheri egg mass in burrow, Merredin (M. Peterson)

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Genus SPICOSPINA

Roberts, Horwitz, Wardell-Johnson, Maxson and Mahony, 1997 Diversity and Evolutionary Relationships A monotypic genus of myobatrachine frogs, discovered in the early 1990s. Restricted to a small area inland of Walpole in southwestern Australia. Molecular studies indicate the sister taxon is Uperoleia, although they diverged long ago (Read et al. 2001; Frost et al. 2006).

Morphological Characteristics Head and body broad, rather flattened; tympanum not visible; eye large with rhomboidal pupil; parotoid glands massive – entire dorsal surface behind head covered with network of elevated, circular glands; ventral skin smooth; fingers and toes elongate and without webbing or lateral fringes. Maxillary teeth present; vomerine teeth absent. Face, hands and feet coloured bright orange in life; dorsum black; venter blue spotted on dark grey to black background.

Reproductive Mode Aquatic reproduction. Eggs laid singly. Tadpoles free-swimming.

Etymology Spicospina refers to the pointed shape of processes on the spine.

Genus SPICOSPINA

Spicospina flammocaerulea Roberts, Horwitz, Wardell-Johnson, Maxson and Mahony, 1997 Sunset Frog

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Walpole (B. Maryan) Rocky Gully (R. Browne-Cooper)

Description. A small to medium-sized frog with massive parotoid glands (2.5 times longer than the eye) and a regular network of circular, raised glands on the back, largest between the eyes, smallest on the flanks. The limbs are short (TL/S-V 0.29–0.32) and the fingers and toes long and lack webbing or fringes. The foot has a small inner metatarsal tubercule but no outer metatarsal tubercle. The back is dark, almost black. The chin and throat back to the insertion of the arms is brilliant orange and the belly is dark grey to black with light blue spots. The hands and feet are brilliant orange and there is an orange patch on either side of the cloaca. Males 26–35 mm; females 31–36 mm S-V.

Distribution. Southwest region. Known only from a small area northeast of Walpole. Habitat. Peaty swamps. Advertisement Call. Two notes rapidly repeated, the first note softer than the second note: ‘da duk…da duk’. Breeding Biology. A late spring to early summer breeder. Males call from October to December from seepages in large hollows or in open water along creek margins. Females lay 50–120 eggs singly among mats of algae. Capsules are opaque (an unusual character) with a size of 3.3–4.1 mm. Tadpoles are small with low-lying fins. Etymology. flammo means ‘red’ and caerulea means ‘blue’ in reference to the colours on the belly. Remarks. The small range of this species coupled with a habitat within that range which is suscepible to fire, has led to this species being listed as ‘threatened and priority fauna’ and ‘fauna that is rare or likely to become extinct’. Further Reading. Calling behaviour – Smith et al. (2003); development – Dziminski and Anstis (2004).

Spicospina flammocaerulea, Walpole (M. Anstis) 117

Genus UPEROLEIA Gray, 1841 Diversity and Evolutionary Relationships A large genus of myobatrachine frogs, with 26 species distributed across most of northern and eastern Australia. In Western Australia, 10 species are found in the Kimberley, 2 in the Pilbara and Gascoyne regions and 1 in the northern central arid zone. The genus was last reviewed in its entirety by Tyler et al. (1981a), and many of the Kimberley species reported here were described in that paper. The sister taxon to Uperoleia is the monotypic Spicospina, but they are only distantly related (Read et al. 2001; Frost et al. 2006).

Morphological Characteristics Head small; body squat; pupil rhomboidal; tympanum not visible; upper jaw with or without teeth; vomerine teeth absent; tongue slender and elongated; dorsal skin usually with large glands (Figure 6A). Limbs very short; fingers unwebbed; first finger shorter than second; toes elongated, cylindrical, with or without webbing; inner metatarsal tubercle not compressed.

Reproductive Mode Aquatic reproduction. Eggs small, pigmented. Where breeding biology is known, spawn typically laid in shallow static water. The larvae are a typical free-swimming tadpole. Males of some species call several meters away from water. Larval life under laboratory conditions took 70 days in one species. However, use of ephemeral water bodies by some species suggests the likelihood of more rapid development in nature.

Etymology Unknown, but probably, like many generic names created by Gray, an arbitrary combination of letters creating a euphonious sound.

Further Reading Call analyses and breeding biology are covered in detail in Tyler et al. (1981a,b,c).

Uperoleia borealis, Kununurra (P. Doughty) 119

FROGS OF WESTERN AUSTRALIA

Uperoleia aspera Tyler, Davies and Martin, 1981 Derby Toadlet

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Broome (G. Gaikhorst)

Description. A moderate-sized species lacking teeth, and possessing moderately developed parotoid and inguinal glands. Skin of back roughened by tubercles. The hindlimbs are short (TL/S-V 0.28–0.34) with slightly webbed toes. Dorsal surface dull brown bearing small and diffuse darker patches. Males 24–35 mm; females 29–34 mm S-V. Distribution. Kimberley region. Dampier Peninsula and southern edge of Yampi Peninsula,

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extending east to Fitzroy Crossing. Habitat. Flooded grasslands. Advertisement Call. A sharp loud click repeated about 3 times a second. Breeding Biology. Presumably similar to other Uperoleia, with aquatic eggs and tadpoles taking 6–8 weeks to complete development. Etymology. aspera means ‘rough’, alluding to the surface of the dorsal skin.

Genus UPEROLEIA

Uperoleia borealis Tyler, Davies and Martin, 1981 Northern Toadlet

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Kununurra (P. Doughty)

Description. A small to moderate-sized species lacking teeth, possessing moderately developed parotoid and inguinal glands. Hindlegs short (TL/S-V 0.29–0.35) with moderate webbing between the toes. The body varies from dull brown to slate, and there are regular-shaped orange patches in the groin and behind the thighs. The throat is slightly stippled with grey. Males 22–32 mm; females 24–32 mm S-V. Distribution. Kimberley region. From Mitchell Plateau south to Mornington Station, and extending through the eastern Kimberley. Also occurs in the adjacent part of the Northern Territory. Habitat. Sparsely vegetated areas of temporarily

inundated grasslands; also occurs along rocky slopes and creeklines. Advertisement Call. A rasping note, with occasional shorter higher-pitched notes emitted. Males call in delayed unison along creeks, creating a ‘wave’ effect. Breeding Biology. Calling males have been found at the base of grass tussocks or in crevices among stones near standing or flowing water. Eggs laid by a captive female had a capsule diameter of 1.8–2.0 mm and an egg diameter of 1.1–1.3 mm. Etymology. borealis means ‘north’ in reference to the distribution of this species.

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FROGS OF WESTERN AUSTRALIA

Uperoleia crassa Tyler, Davies and Martin, 1981 Fat Toadlet

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Mornington Station (J. Francis)

Description. A small to medium-sized species with a rotund but roughly oval body shape. It lacks teeth and has short limbs (TL/S-V 0.29– 0.35), with only a trace of webbing between the toes. The back and sides of the body are a uniform dark brown, usually with gold markings upon the parotoid glands. The flanks and sides of the head are pale slate, and the undersurface of the body is dull cream tinged with grey between the jaws. Males 24–31 mm; females 30–34 mm S-V. Distribution. Kimberley region. The northwest portion including Truscott, Mitchell Plateau,

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Prince Regent River Nature Reserve, Mornington Station and Drysdale River Station. Habitat. Flooded grasslands with sparse Livistona and Eucalyptus forest. Advertisement Call. A slow rasp with several pulses. Breeding Biology. Males call from the base of grass tussocks close to water in temporarily flooded areas. Females can lay more than 350 eggs. Etymology. crassa means ‘fat’ in reference to the round shape of this species.

Genus UPEROLEIA

Uperoleia glandulosa Davies, Mahony and Roberts, 1985 Glandular Toadlet !! !!!! !

Strelly River (B. Maryan)

Description. A small Uperoleia which lacks teeth. Snout short and truncate when viewed from above and in profile. The limbs are moderate (TS/S-V 0.34–0.41) and the dorsal glands conspicuous. There is a large inner and a broad outer metatarsal tubercle and webbing only at the base of the toes. Ventral surface finely granular. The back is olive-brown with black spots and blotches, an anterior indistinct series of vertebral orange spots and a broken orange dorsolateral stripe from behind the orbit to the groin. There is also a small orange spot on each side of the sacrum and a large orange-red blotch in the groin and on the outer surface of the thigh.

Males 20–31 mm; female 25 mm S-V. Distribution. Arid zone. Occurs in the northern Pilbara from the Strelly River south to Woodstock. Habitat. Found along creeks and low-lying areas subject to seasonal flooding. Advertisement Call. A sharp single-noted ‘click’. Breeding Biology. Davies et al. (1985) found males at the base of sedge clumps close to the water in roadside drains after heavy rain. Etymology. glandulosa refers to the conspicuous glands of this species.

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FROGS OF WESTERN AUSTRALIA !!

Uperoleia lithomoda Tyler, Davies and Martin, 1981 Stonemason Toadlet

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Kununurra (B. Maryan)

Description. A small robust, short-legged species (TL/S-V 0.31–0.36) lacking teeth, and with only a trace or no webbing between the toes. The parotoid and inguinal glands are moderately developed and usually connect along the sides. The back ranges from a dull tan colour to brown, and most frogs have a narrow, cream stripe dividing the head. The undersurface of the body is pale cream, and there may be large areas of dark grey upon the chest. Leg patches are red. The throat of the males is blackish. Males 21–27 mm; females 22–32 mm S-V. Distribution. Kimberley region. South to Napier Downs, Mornington Station and east to Lake

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Argyle. Also in tropical Northern Territory, Queensland and New Guinea. Habitat. Open tussock grassland in areas exposed to flooding. Advertisement Call. An explosive ‘tick’, resembling a stone being chipped. Breeding Biology. Males call from exposed positions on the ground up to 10 m from water. Etymology. lithomoda means ‘stonemason’ in reference to the call. Further Reading. Call analysis and tadpoles – Tyler et al. (1981a,c), Davies et al. (1986).

Genus UPEROLEIA

Uperoleia marmorata Gray, 1841 Marbled Toadlet

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Holotype (C. McCarthy/BMNH)

Description. This small species is poorly documented. The only specimen known is a male of 30 mm S-V. It possesses maxillary teeth, short limbs (TL/S-V 0.35) and has a trace of webbing between the toes. Gray (1841) described its colour as ‘black and green marbled leaving a triangular greenish spot on the forehead, beneath lead-colour’. Distribution. Kimberley region. Known only from the original collection site which, according to maps published by J. E. Grey (1841), might be near the mouth of the Prince Regent River. Habitat. Unknown.

Advertisement Call and Breeding Biology. Unknown. Etymology. marmorata means ‘marbled’ in reference to its appearance. Remarks. This is the type species for the genus Uperoleia. Before the revision of Tyler et al. (1981a), its range was erroneously believed to extend from the Kimberley across northern Australia and into New South Wales. Now its distribution is confined to a location that is not known precisely.

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FROGS OF WESTERN AUSTRALIA

Uperoleia micra Doughty and Roberts, 2008 Tiny Toadlet

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Katers Island (P. Doughty)

Description. A small species with tubercular skin, short limbs (TL/S-V 0.34–0.38) and basal webbing between the toes. The back is dark brown to black with darker blotches discernible. The parotoid glands, upper arms and snout often have a reddish hue, and the loreal and lateral zones are stippled with pale bluish-white spots; belly and chin speckled. Males 18–22 mm; female 21 mm S-V. Distribution. Kimberley region. Known from

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the Prince Regent River, Walcott Inlet, Katers Island and Boongaree Island. Habitat. Associated with sandstone outcrops. Advertisement Call. A high-pitched rasp. Breeding Biology. Males were observed calling from boulder piles with flowing water or on sandstone platforms with slow flowing water and rock pools among Triodia clumps. Etymology. micra means ‘small’ in reference to the body size.

Genus UPEROLEIA

Uperoleia micromeles Tyler, Davies and Martin, 1981 Tanami Toadlet ! !

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Telfer (G. Harold)

Description. A small squat species with a broad snout (the only species of Uperoleia in which the internarial distance is greater than the distance between the eye and the nares). The limbs are very short (TL/S-V 0.28–0.33). The back is smooth, and the toes (the fifth of which is very narrow) are unwebbed or nearly so. The metatarsal tubercles are very large. The back is a rich dull brown and the parotoid glands suffused with metallic gold. Undersurface immaculate.

Male 24 mm; females 24–31 mm S-V. Distribution. Northeastern arid zone. Found at scattered localities in the Tanami and the Great Sandy Desert. Habitat. Low dunes with Triodia and soft grasses. Advertisement Call and Breeding Biology. Unknown. Etymology. micromeles means ‘small-limbed’.

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FROGS OF WESTERN AUSTRALIA

Uperoleia minima Tyler, Davies and Martin, 1981 Small Toadlet

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Mitchell Plateau (P. Doughty)

Description. A very small species with moderate legs (TL/S-V 0.30–0.39), long unwebbed toes and a very coarsely tubercular dorsal skin. There are no maxillary teeth. The back is pale brown and bears slightly darker and indistinct streaks. There are broad, straw-coloured bands on the upper forearms and patches of scarlet in the groin and behind the thighs. The undersurface of the body is white, with indistinct patches of pale grey on the throat, chest and flanks. Males 16–23 mm S-V.

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Distribution. Kimberley region. Known only from the Mitchell Plateau. Habitat. Dense grasslands subject to flooding. Advertisement Call. A single sharp click produced from the base of grass tussocks. Breeding Biology. Unknown. Etymology. minima means ‘smallest’ in reference to this species’ tiny adult size.

Genus UPEROLEIA

Uperoleia mjobergii (Andersson, 1913) West Kimberley Toadlet

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King Leopold Ranges (G. Gaikhorst)

Description. A small and distinctive species. It possesses maxillary teeth, short hindlimbs (TL/ S-V 0.28–0.36) and a trace of webbing between the toes. There is a small but prominent papilla upon the heel, and the glands upon the flanks are vast, usually joining to the back of the parotoid glands. The species has a pale background colour with large dark olive or brown blotches. The dorsal glands are a pale yellowish-brown. Males 19–25 mm; females 21–23 mm S-V. Distribution. Kimberley region. Dampier Peninsula east to Fitzroy Crossing.

Habitat. Open floodplains and grasslands. Hides beneath debris on the surface. Advertisement Call. A short rasp. Breeding Biology. Breeds following heavy rains. Spawn and tadpoles unknown. Etymology. Named for Eric Georg Mjöberg (1882–1938), leader of the Swedish expedition to northern Australia that led to the description of this species. Further Reading. Redescription – Tyler et al. (1981a); call analysis – Tyler et al. (1981b).

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FROGS OF WESTERN AUSTRALIA

Uperoleia russelli (Loveridge, 1933) Northwest Toadlet ! ! !! !!! !! ! ! ! ! ! ! ! ! !! ! ! !! ! !!! ! ! !! ! !!! ! !!! ! ! ! !! !!!

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♁ Millstream-Chichester NP (B. Maryan) ♂ Mt. Brockman (P. Doughty)

Description. A medium-sized robust species which lacks teeth, has conspicuous parotoid and inguinal glands, short hindlimbs (TL/S-V 0.29– 0.35) and extensive webbing between the toes. A dull reddish-brown with discrete dark blotches and often a thin vertebral stripe. The parotoid and coccygeal glands are reddish. Males 27–35 mm; females 28–39 mm S-V.

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Distribution. Arid zone. The Pilbara and south to Shark Bay. Habitat. Most abundant in low-lying areas, especially creek beds. Advertisement Call. A short rasp with occasional shorter high-pitched calls given. Breeding Biology. Main and Storr (1966) found amplexed pairs in shallow, slow-moving water or on nearby patches of silt. Etymology. Named for A. R. E. Russell who owned the property where the first specimens were collected. Further Reading. Redescription – Tyler et al. (1981a); diet – Main and Calaby (1957).

Genus UPEROLEIA

Uperoleia talpa Tyler, Davies and Martin, 1981 Mole Toadlet

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Derby (D. Moore)

Description. A large and very robust species that lacks teeth and has short legs (TL/S-V 0.29– 0.31) with well developed webbing between the toes. The parotoid and inguinal glands are moderately to well developed. The back is bronze with darker markings; the parotoid glands have a distinct gold lustre. Throat black; chest stippled with black. Abdomen pearl and undersurface of thighs creamish. Males 26–40 mm; females 35–42 mm S-V. Distribution. Kimberley region. Dampier Peninsula and east to Fitzroy Crossing.

Habitat. Sparsely vegetated plains. Advertisement Call. A plaintive ‘yelp’. Breeding Biology. Males call from leaf litter on the margins of flooded grasslands. Larval lifespan is about 2 months. Etymology. talpa means ‘mole’ for this species’ appearance and subterranean habits. Further Reading. Redescription and call analysis – Davies and Martin (1988); tadpoles – Davies and Watson (1998).

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FROGS OF WESTERN AUSTRALIA

Uperoleia trachyderma Tyler, Davies and Martin, 1981 Blacksoil Toadlet

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Fitzroy Crossing (D. Moore)

Description. A relatively flattened species with a dense covering of small conical tubercles on the back. The legs are short (TL/S-V 0.35), the toes are long and unwebbed and the metatarsal tubercles poorly developed. There are no teeth. The back is dull slate with obscure, slightly darker markings, the dorsal glands are sometimes bright yellowish cream and the tubercles orange. The thigh patches are vivid orange red and nearly connect on top of the thigh. Throat and undersurface of body white, pigmented with

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grey. Males 17–28 mm S-V; female 24 mm S-V. Distribution. Kimberley region. West to Fitzroy Crossing and east to Kununurra. Also occurs in the Northern Territory. Advertisement Call. A short creak. Breeding Biology. Males call away from water in near shallow flooded grassy areas on blacksoils or sandy soils. Etymology. trachyderma means ‘rough-skinned’ in reference to the conical tubercles on the back.

Genus UPEROLEIA

Uperoleia aspera

Uperoleia borealis

Uperoleia crassa

Uperoleia lithomoda

Uperoleia micra

Uperoleia minima

Uperoleia mjobergii

Uperoleia talpa

Sonograms of Uperoleia species from the Kimberley region. The time interval of all calls is 1 second along the X-axis. Calls that appear as ‘spikes’ sound like sharp clicks or taps to the ear; calls that appear as several or many pulses sound like rasps or squelches. Calls of all species can be heard on the Alcoa Frog Watch web site – http://frogwatch.museum.wa.gov.au). 133

Genus Bufo Laurenti, 1768 Diversity and Evolutionary Relationships Bufo largely refers to amphibians commonly referred to as ‘toads’. Toads, however, are true frogs that have evolved a suite of characters (rough dark skin, terrestrial habits) that people conviently classify as ‘toads’. They are a cosmopolitan group, except naturally not occurring in Australia, New Zealand, New Guinea and South Pacific Islands, with the nearest native species occurring in Indonesia (e.g. B. melanostictus). Recently, there have been several conflicting proposals to recognize multiple genera within Bufo (Frost et al. 2006; Brandáo et al. 2007; Pramuk et al. 2008). The names Chaunus Wagler, 1828 or Rhinella Fitzinger, 1826 may apply to B. marinus in Australia. We adopt the more traditional concept of Bufo here pending taxonomic stability.

Morphological Characteristics A diverse group of frogs that range from 20–250 mm S-V. They usually have thick glandular skin and heavily ossified skulls, often coosified with the skin giving their faces a bony appearance. Most species tending towards terrestrial habits and have dark rough skin and short limbs.

Reproductive Mode Large masses of eggs in long strings are laid in shallow water in static or slow-flowing water. Amplexus is axillary.

Etymology Bufo is Latin for ‘toad’.

Genus Bufo

Bufo marinus Linnaeus, 1758 Cane Toad

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Fogg Dam, NT (P. Doughty) Juvenile : Bradshaw, NT (J. Francis)

Description. A very large robust species with diagnostic bony ridges around the eye and snout (not shared by any native frog). Head is wider than long, and the face is short. The parotoid glands are very large and prominent. The limbs are moderately short (TL/S-V 0.35–0.43); the fingers are unwebbed with moderately extensive webbing between the toes. The resting posture is more upright than in native species. The skin of males is coarse; that of females less so. Colour is yellow-brown; the ventral surfaces are white, with some blotching in males. Males 80–150 mm; females 90–240 mm S-V. Distribution. At this writing (early 2009), the first Cane Toads have crossed the (WA-NT) border and are poised to invade the eastern Kimberley. In Australia occuring throughout

tropical Northern Territory and Queensland, and extending south to Port Macquarie in New South Wales. Habitat. Occurs in most environments, but prefers open grasslands. Advertisement Call. A long steady trill. Breeding Biology. A summer breeder. Large females can lay clutches of 30,000 or more eggs. Breeding occurs in shallow water bodies with little fringing vegetation and a shallow slope. Tadpoles are small (to 27 mm), jet black (including the belly) and form large schools. Metamorphosis takes only 1 month and metamorphs are tiny (5–10 mm)(Anstis 2002). All eggs, tadpoles and metamorphs are toxic. Etymology. marinus means ‘marine’. Further Reading. The cane toad is one of the world’s most well-studied amphibians. Tyler (1994) provides a general overview including their history of introduction in Australia, whereas Lever (2001) is the most comprehensive and detailed. Further information and the results of recent research on the impact of cane toads on native fauna can be found at www.canetoadsinoz. com, a web site maintained by the laboratory of Rick Shine (University of Sydney).

Bufo marinus, Bradshaw (J. Francis) 135

CHECKLISTS OF FROGS FOR URBAN CENTRES Checklists contain species that occur within a 50 km radius of the town or city. SOUTHWEST REGION Geraldton Family Hylidae

Litoria adelaidensis Litoria moorei Family Limnodynastidae



Heleioporus albopunctatus Heleioporus eyrei Heleioporus psammophilus Limnodynastes dorsalis Neobatrachus pelobatoides Neobatrachus wilsmorei

Family Myobatrachidae

Crinia pseudinsignifera Myobatrachus gouldii Pseudophryne guentheri Perth Family Hylidae

Litoria adelaidensis Litoria moorei Family Limnodynastidae



Heleioporus albopunctatus Heleioporus barycragus Heleioporus eyrei Heleioporus inornatus Heleioporus psammophilus Limnodynastes dorsalis Neobatrachus pelobatoides

Family Myobatrachidae



Crinia georgiana Crinia glauerti Crinia insignifera Crinia pseudinsignifera Geocrinia leai Myobatrachus gouldii Pseudophryne guentheri

Bunbury Family Hylidae

Litoria adelaidensis Litoria moorei Family Limnodynastidae

Heleioporus eyrei Heleioporus inornatus 136

Heleioporus psammophilus Limnodynastes dorsalis Family Myobatrachidae



Crinia georgiana Crinia glauerti Crinia insignifera Crinia pseudinsignifera Geocrinia leai Pseudophryne guentheri

Albany Family Hylidae

Litoria adelaidensis Litoria cyclorhyncha Litoria moorei Family Limnodynastidae



Heleioporus eyrei Heleioporus inornatus Heleioporus psammophilus Limnodynastes dorsalis

Family Myobatrachidae



Crinia georgiana Crinia glauerti Crinia pseudinsignifera Crinia subinsignifera Geocrinia leai Metacrinia nichollsi Pseudophryne guentheri

Esperance Family Hylidae

Litoria adelaidensis Litoria cyclorhyncha Family Limnodynastidae



Heleioporus eyrei Limnodynastes dorsalis Neobatrachus albipes Neobatrachus pelobatoides

Family Myobatrachidae



Crinia georgiana Crinia pseudinsignifera Crinia subinsignifera Myobatrachus gouldii Pseudophryne guentheri

ARID ZONE Port Hedland Family Hylidae

Cyclorana australis Cyclorana maini Litoria rubella

CHECKLISTS OF FROGS FOR URBAN CENTRES

Family Limnodynastidae

Neobatrachus aquilonius Notaden nichollsi Platyplectrum spenceri Family Myobatrachidae

Uperoleia glandulosa Uperoleia russelli Exmouth Family Hylidae

Cyclorana maini Litoria rubella Family Limnodynastidae

Neobatrachus fulvus Neobatrachus sutor Platyplectrum spenceri Family Myobatrachidae

Pseudophryne douglasi



Litoria nasuta Litoria pallida Litoria rothii Litoria rubella Litoria splendida Litoria staccato Litoria watjulumensis

Family Limnodynastidae



Limnodynastes convexiusculus Limnodynastes depressus Limnodynastes lignarius Notaden melanoscaphus Platyplectrum ornatum

Family Myobatrachidae



Crinia bilingua Uperoleia borealis Uperoleia lithomoda Uperoleia trachyderma

Newman

Derby

Family Hylidae

Family Hylidae

Cyclorana maini Cyclorana platycephala Litoria rubella Family Limnodynastidae



Neobatrachus sutor Neobatrachus wilsmorei Notaden nichollsi Platyplectrum spenceri

Family Myobatrachidae

Uperoleia russelli Kalgoorlie Family Limnodynastidae

Neobatrachus kunapalari Neobatrachus sutor Neobatrachus wilsmorei Family Myobatrachidae

Pseudophryne occidentalis KIMBERLEY REGION Kununurra Family Hylidae



Cyclorana australis Cyclorana cryptotis Cyclorana cultripes Cyclorana longipes Cyclorana vagitus Litoria bicolor Litoria caerulea Litoria coplandi Litoria inermis Litoria meiriana



Cyclorana australis Cyclorana cryptotis Cyclorana longipes Cyclorana vagitus Litoria caerulea Litoria inermis Litoria nasuta Litoria pallida Litoria rubella

Family Limnodynastidae

Neobatrachus aquilonius Notaden nichollsi Platyplectrum ornatum Family Myobatrachidae

Uperoleia aspera Uperoleia mjobergii Uperoleia talpa Broome Family Hylidae



Cyclorana australis Cyclorana longipes Litoria caerulea Litoria nasuta Litoria rothii Litoria rubella

Family Limnodynastidae

Notaden nichollsi Family Myobatrachidae

Uperoleia aspera Uperoleia mjobergii Uperoleia talpa 137

GLOSSARY Annulus: narrow, ring-like structure surrounding the tympanum. Arboreal: adapted to life above the ground; climbing. Axillary amplexus: mating embrace of frogs, with the male grasping the female behind her arms. Biogeography: study of the distribution of animals and plants. Choanae: paired openings on the palate communicating with the nostrils. Cloaca: common opening of urinary, digestive and reproductive tracts. Colloidal: a fine suspension in water producing a dense, milky or opaque appearance. Complex call: when referring to a male advertisement call, means two or more different parts. Cordiform: shaped in the form of a heart. Dextral: on the right side; to the right of the midline. Diurnal: active in the daytime. Dorsal: on the upper surface or back. Dorsolateral: dividing the upper surface (dorsal) from the side (lateral). Endemic: native to a particular area. Eustachian tubes: canals connecting the ear to the back portion of the mouth cavity. Fossorial: adapted for life below the ground; digging or burrowing. Herpetology: the study of amphibians and reptiles. Inguinal: in the region of the groin, referring to the junction of the posterior end of the side of the body with the thigh. Inguinal amplexus: mating embrace of frogs, the male grasping the female around the abdomen, anterior to her legs. Iris: the coloured zone surrounding the pupil of the eye. Keratin: a hard protective layer secreted by the outer cells of the skin, and forming in male frogs the nuptial pads on the fingers. Labial: of the lips; describing the superficial rows of teeth of tadpoles. Larval: of the larva (tadpole). Lateral: on the side or flank. Lentic: static water such as a pond or swamp. Lore: the area between the nostril and eye. Lotic: flowing water. Lyrate: with a pair of bow-like curves facing one another. Macroglands: enlarged skin glands. Mandibular: of the lower jaw. Maxillary: of the upper jaw. Medial: of the middle. Metamorphosis: the change of state taking place in an animal’s life-history; e.g. from tadpole to frog. 138

GLOSSARY

Metatarsus: longest bone in the sole of the foot. Morphology: the study of form and structure. Mid-vertebral: a line running down the centre of the back, above the vertebral column. Monotypic: a genus containing only one species. Nares: external openings of the nostrils. Nomenclature: the naming of animals, plants or things. Nocturnal: active at night. Nuptial Pads: darkened pads (or spikes) on the first (and sometimes second) fingers of male frogs, used to grasp the female during amplexus. Palatine: a bone of the palate or the roof of the mouth. Palmar: of the palm of the hand. Papillae: small, raised structures surrounding the mouth of tadpoles. Also used to describe small structures found on the heels of some species of frogs. Parotoid: large glands behind the head of some frogs; often contain toxic substances. Pupil: the central ‘window’ of the eye. Rhomboid: shaped like a diamond. Rostral: on the snout. Serrated: having a toothed or saw-like edge. Sexual dimorphism: having two distinguishable forms or kinds, e.g. difference in size, structure or colour of male and female. Spawn: eggs surrounded by jelly; generally applied to groups of eggs. Spiracle: external aperture draining water from the mouth of a tadpole. Subarticular: beneath a joint between two bones, usually in reference to fingers or toes. Subterminal: below one end; e.g. a subterminal mouth is slightly behind the normal position. Being beneath rather than at the end of the head. Supralabial: above the upper lip. Supratympanic: above the tympanum. Terrestrial: adapted to life upon the ground. Tibia: bone of the lower limb extending from the knee to the ankle. Transocular: the upper surface of the head between the eyes; commonly used to describe a bar of colour uniting the eyelids. Tubercle: a raised structure upon the skin; commonly found beneath joints and upon the palm and sole. Tubercular: bearing tubercles. Tympanum: external ear; surrounded by an annulus. Ventral: of the undersurface or belly. Vomerine: paired bones on the palate, commonly bearing a row of teeth.

139

MAP OF REGIONAL DIVISIONS

KIMBERLEY (summer rainfall)

ARID ZONE (low irregular rainfall)

SOUTH WEST (winter rainfall)

140

Key To Adult Frogs Of The Southwest Region 1

Discs present on tips of fingers and toes (Figure 11); belly granular (Figure 5) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 (Hylidae) Tips of fingers and toes lack discs; belly smooth or granular. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 2

White stripe on upper lip, continuing along side of body . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Litoria adelaidensis (p. 68) Upper lip and side of body lack white stripe . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 3

Large yellow, oval or circular spots in groin and thigh (Figure 22) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Litoria cyclorhyncha (p. 73) Groin and thighs without spots. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Litoria moorei (p. 78) 4

Large compressed, shovel-shaped inner metatarsal tubercle present on sole of foot; medium-large size (Figure 9) . . . . . . . . . . . . . . . . . . 5 (Limnodynastidae) No large compressed, shovel-shaped metatarsal tubercle on foot (small, rounded metatarsal tubercle may be present); small-medium size . . . . . . . . . . . . . . . . . . . . . . . . 15 (Myobatrachidae) 5

Large oval gland on upper surface of calf; vomerine teeth in row across palate behind choanae . . . . . . . . . . . . Limnodynastes dorsalis (p. 64) No gland on upper surface of calf; vomerine teeth in short series between choanae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 6

Pair of diverging yellow or cream lines commencing behind each eye; one extending along flank, the other converging on vertebral stripe, forming a ‘W’ mark . . . . . . . . . . . . . . . . . . . . Neobatrachus wilsmorei (p. 101) No diverging lines behind eye . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 7

Numerous large cream or white spots on back and sides of body. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Heleioporus albopunctatus (p. 54) No large cream or white spots on back and sides of body. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 8

Back uniform chocolate or coppery brown . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 Back marbled or blotched grey, brown or yellow . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 9

Zone of yellow blotches and tubercles on flanks sharply differentiated from plain chocolate-brown back. . . . . . . . . . . . . . . . Heleioporus barycragus (p. 55) No zone of yellow blotches or spots on flanks. Back and flanks coppery-brown . . . . . . . . . . . . . . . . . . . . . . . . . . . . Heleioporus inornatus (p. 57) 10

Back rough, greenish-yellow with grey marbling; fine red vertebral stripe usually present . . . . . . . . . . . . . . . . . . . Neobatrachus pelobatoides (p. 99) 141

FROGS OF WESTERN AUSTRALIA

Back smooth, predominantly brown, gold or grey; fine red vertebral stripe absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11 11

Area of parotoid gland usually with a distinct yellow wash beneath brownish pattern; call is a long, single moan . . . . . . . . . . . . . . . Heleioporus eyrei (p. 56) No yellow colour on parotoid glands; call not a single low moan . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12 12

Flanks pale brown with white mottling; tympanum indistinct . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Heleioporus psammophilus (p. 58) Flanks not pale brown with white mottling; tympanum distinct . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13 13

Inner metatarsal tubercle weakly pigmented (Figure 23); upper surface of foot white . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Neobatrachus albipes (p. 94) Inner metatarsal tubercle black or brown; upper surface of foot pale brown . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14 14

Call a high pitched trill; moderate webbing between toes; males with small black spines on back. . . . . . . . . . . . . . . . . . . . . . Neobatrachus kunapalari (p. 98) Call a repetitive tapping; extensive webbing between toes; no spines on males. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Neobatrachus sutor (p. 100) 15

Chin, throat and chest brilliant orange; belly grey-black with light blue spots. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Spicospina flammocaerulea (p. 117) Chin, throat and chest not brilliant orange; belly not grey-black and lacking light blue spots. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16 16

Red patches in groin and on thighs; eyelid golden . . . . . . . . . . . . . . . . . . Crinia georgiana (p. 33) No red patches in groin, or on thighs; eyelid not golden. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17 17

Belly immaculate, limbs short, body turtle-like, head and eyes small. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Myobatrachus gouldii (p. 91) Belly freckled or blotched grey, black or brown or suffused with pink or yellow. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18 18

Fine, pale vertebral stripe usually present; metatarsal tubercles barely discernible. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19 Fine pale vertebral stripe rarely present; at least one rounded metatarsal tubercle prominent. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20 19

Eyes slightly protruding, pale background colour . . . . . . . . . . . . . . . Arenophryne rotunda (p. 26) Eyes small and not protruding, cream to brown background colour. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Arenophryne xiphorhyncha (p. 27) 142

KEY TO THE SOUTHWEST REGION

20

Frogs walk; upper jaw not toothed.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21 Frogs hop; upper jaw toothed.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23 21

Distinct orange markings at insertion of limbs on ventral surface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Metacrinia nichollsi (p. 89) No orange markings on ventral surface. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22 22

Heel with two tubercles of equal size (Figure 26). . . . . . . . . . . . . Pseudophryne guentheri (p. 113) Heel with two tubercles of unequal size (the outer the smaller; Figure 27). . . . . . . . . . . . . . Pseudophryne occidentalis (p. 114) 23

Belly smooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24 (Geocrinia) Belly granular . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28 (remaining Crinia) 24

Belly greenish-yellow; tips of toes slightly dilated. . . . . . . . . . . . . . . . . . . . . Geocrinia leai (p. 49) Belly not greenish-yellow; tips of toes not dilated. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25 25

Belly pink . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Geocrinia rosea (p. 51) Belly not pink . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26 26

Belly bright orange or yellow. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Geocrinia vitellina (p. 52) Belly not bright orange or yellow . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27 27

Belly white or with yellowish wash . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Geocrinia alba (p. 48) Belly fawn-brown. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Geocrinia lutea (p. 50) 28

Call resembles a squelch . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29 Call does not resemble a squelch . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30 29

Flanges on toes narrow; species restricted to western coastal plain . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Crinia insignifera (p. 36) Flanges on toes moderately expanded; species restricted to the south coast. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Crinia subinsignifera (p. 38) 30

Call is a wavering bleat, throat of male not entirely black (Figure 20A) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Crinia pseudinsignifera (p. 37) Call a rattling or clicking sound, throat of male entirely black . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Crinia glauerti (p. 35)

143

Key to adult frogs of the Arid zone 1

Fingers and toes with terminal discs (Figure 11) . . . . . . . . . . . . . . . . . . . . . Litoria rubella (p. 82) Fingers and toes without terminal discs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 2

Large, compressed, shovel-shaped inner metatarsal tubercle present (Figure 9) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 Large, compressed, shovel-shaped inner metatarsal tubercle absent (small, rounded metatarsal tubercles may be present) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15 3

Upper jaw not toothed; frogs may release copious amounts of adhesive secretion when handled . . . . . . . . . . . . . . . . . . . . . Notaden nichollsi (p. 105) Upper jaw toothed; frogs never release copious amounts of adhesive secretion when handled . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 4

Vomerine teeth (Figure 12) in long row behind choanae . . . . . . . . Platyplectrum spenceri (p. 109) Vomerine teeth in short row between choanae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 5

Pair of diverging yellow or cream lines commencing behind each eye; one extending along flank, the other converging on vertebral stripe so forming a ‘W’. . . . . . . . . . . . . . Neobatrachus wilsmorei (p. 101) No diverging yellow or cream lines. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 6

Prominent dark streak through lore and tympanum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 No dark streak through lore and tympanum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 7

Dorsolateral fold present (Figure 7) . . . . . . . . . . . . . . . . . . . . . . . . . . . Cyclorana australis (p. 40) Dorsolateral fold absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 8

Dull grey or brown with poorly-contrasting irregular dark blotches; moderately extensive webbing between toes. . . . . . . . . . . . Cyclorana maini (p. 44) Highly contrasting olive blotches on yellow-brown background; basal webbing between toes . . . . . . . . . . . . . . . . . . . . . . . . Cyclorana longipes (p. 43) 9

Back grey or tan, marbled darker grey or green; toes fully webbed (Figure 10) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cyclorana platycephala (p. 45) Back marbled gold and brown or gold and blackish-brown; toes partially webbed. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 10

Inner metatarsal tubercle weakly pigmented. . . . . . . . . . . . . . . . . . . . Neobatrachus albipes (p. 94) Inner metatarsal tubercle black or brown . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11

144

KEY TO THE ARID ZONE

11

Call a repetitive tapping which does not carry a long way. . . . . . . . . . Neobatrachus sutor (p. 100) Call a high pitched trill which carries a long way. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12 12

Back chocolate brown with yellow or gold variegations . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13 Back light brown, yellowish or bronze . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14 13

Back chocolate-brown with yellow mottling and usually vertebral stripe; body size large (48–59 mm S-V); found in southern Kimberley, coastal Pilbara and arid interior. . . . . . . . . . Neobatrachus aquilonius (p. 95) Back chocolate brown with gold variegations; no vertebral stripe; medium (38–48 mm S-V); only found between Northwest Cape and Shark Bay. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Neobatrachus fulvus (p. 97) 14

Medium-sized species (41–49 mm) . . . . . . . . . . . . . . . . . . . . . . . . . Neobatrachus centralis (p. 96) Large species (48–61 mm). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Neobatrachus kunapalari (p. 98) 15

Fifth toe much narrower than other toes . . . . . . . . . . . . . . . . . . . . . Uperoleia micromeles (p. 127) Toes equal in thickness . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16 16

Toes partially webbed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17 Toes unwebbed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18 17

Call a sharp click; toes basally webbed . . . . . . . . . . . . . . . . . . . . . . . Uperoleia glandulosa (p. 123) Call a short rasp; toes extensively webbed. . . . . . . . . . . . . . . . . . . . . . . . Uperoleia russelli (p. 130) 18

Belly granular . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pseudophryne douglasi (p. 112) Belly smooth (throat and groin may be granular). . . . . . . . . . . . Pseudophryne occidentalis (p. 114)

145

Key to adult frogs of the Kimberley region 1

Bony ridges around eye and nostrils; parotoid glands massive; large body size (to 24 cm S-V) . . . . . . . . . . . . . . . . . . . . . . . Bufo marinus (p. 135) No bony ridge on face; parotoid glands not massive; body size < 12 cm S-V. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 2

Tympanum large and conspicuous; its diameter equal to the diameter of the eye; male nuptial pad a cluster of spines . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Limnodynastes lignarius (p. 65) Tympanum visible or not visible, if visible its diameter less than the eye diameter; nuptial pad not in the form of large spines . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 3

Continuous elongated skin folds along back . . . . . . . . . . . . . . . . . . . . . . . . Litoria nasuta (p. 79) Elongated skin folds absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 4

Tips of fingers and toes with prominent terminal discs (Figure 11) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 Tips of fingers and toes without prominent terminal discs. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15 5

Solid or diffuse stripe through lore to tympanum, sometimes extending on to body . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 No stripe through lore to tympanum, or on body . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 6

Toes webbed to discs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 Toes not webbed to discs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 7

Loreal stripe bounded by a white stripe below; small species (to 35 mm S-V) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Litoria bicolor (p. 69) Loreal stripe not bounded by white stripe below; medium-large species. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Litoria watjulumensis (p. 86) 8

Head short; fingers webbed at base. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Litoria rubella (p. 82) Head elongated; fingers not webbed. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 9

Brown loreal and side stripe bounded by white below; no stripes on back; very small species (to 17 mm S-V) . . . . . . . . . . . . . . Litoria microbelos (p. 77) Diffuse loreal stripe not bounded by white below; diffuse dorsolateral stripes often present; medium-sized species (to 40 mm S-V) . . . . . . . . . . . . . . . . . . . . . . . . . . . . Litoria staccato (p. 84) 10

Large protruding gland on top of head; head and body usually with pale yellow spots . . . . . . . . . . . . . . . . . . . . . Litoria splendida (p. 83) 146

KEY TO THE KIMBERLEY REGION

No protruding gland on top of head; no yellow spots on head or body . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11 11

Upper segment of iris red; supra-tympanic fold edged with black. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Litoria rothii (p. 81) Upper segment of iris not red; supratympanic fold (if present) not edged with black . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12 12

Back green, olive or greenish-brown without pattern; back of thighs not mottled . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13 Back and legs slate or brown; back usually mottled, back of thighs always mottled, to 42 mm S-V. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14 13

Robust body (to 110 mm S-V); green or olive; skin smooth. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Litoria caerulea (p. 70) Flattened body (to 60 mm S-V); greenish-brown; skin granular. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Litoria cavernicola (p. 71) 14

Back and thighs slate or dark brown with cream or yellow spots or blotches on back of thighs (to 21 mm S-V); diurnal and nocturnal. . . . . . . . . . . . . . . . . . . . . . . . . . . Litoria meiriana (p. 76) Back pale brown; back of thighs dark brown with cream spots or flecks (to 42 mm S-V); nocturnal. . . . . . . . . . . . . . . . . . . . Litoria coplandi (p. 72) 15

Large to moderately large compressed shovel-shaped inner metatarsal tubercle present; outer metatarsal tubercle, if present, poorly developed (Figure 9). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16 Large to moderately large compressed shovel-shaped inner metatarsal tubercle absent; small rounded metatarsal tubercles may be present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26 16

Upper jaw not toothed; frogs may release copious amounts of adhesive secretion when handled . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17 Upper jaw toothed; frogs never release large amounts of adhesive secretions when handled . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19 17

Inner metatarsal tubercle black . . . . . . . . . . . . . . . . . . . . . . . . . . . Notaden melanoscaphus (p. 104) Inner metatarsal tubercle unpigmented . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18 18

Black tubercles on back . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Notaden nichollsi (p. 105) Orange tubercles on back . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Notaden weigeli (p. 106) 19

Vomerine teeth (Figure 12) in row behind choanae, foamy egg mass. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20 Vomerine teeth in row between choanae, non-foamy egg mass . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21 147

FROGS OF WESTERN AUSTRALIA

20

Back with dark, slightly raised tubercles; body shape flattened. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Limnodynastes convexiusculus (p. 62) Back smooth; body round . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Platyplectrum ornatum (p. 108) 21

Belly smooth; pupil vertical; inner metatarsal tubercle brownish . . . . . . . . . . . . . . . . . . . . . . . . . . . . Neobatrachus aquilonius (p. 95) Belly granular (Figure 5); pupil not vertical; inner metatarsal tubercles unpigmented. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22 22

Tympanum absent (Figure 3) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cyclorana cryptotis (p. 41) Tympanum present. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23 23

Conspicuous dorsolateral fold on body (Figure 7). . . . . . . . . . . . . . . . . Cyclorana australis (p. 40) No dorsolateral fold on body . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24 24

Dark markings on back sharply distinguished from pale background; pale transocular bar prominent. . . . . . . . . . . . . . . . . . Cyclorana longipes (p. 43) Markings on back dull and not readily distinguished from background colour . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25 25

Call a short harsh note; to 43 mm S-V; east Kimberley . . . . . . . . . . . . . Cyclorana cultripes (p. 42) Call a wailing; to 48 mm S-V; southern Kimberley . . . . . . . . . . . . . . . . Cyclorana vagitus (p. 46) 26

Toes webbed to tips; back greenish or slate. . . . . . . . . . . . . . . . . . . . . . . . . . . Litoria dahlii (p. 74) Toes not webbed to tips; back not greenish . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27 27

Tympanum distinct . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28 Tympanum indistinct . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31 28

Toes not webbed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Limnodynastes depressus (p. 63) Toes partly webbed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29 29

Upper edge of tibia with a continuous dark brown stripe . . . . . . . . . . . . . . . Litoria tornieri (p. 85) Tibia with or without dark markings on upper margins; if markings present, not continuous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30 30

Skin of back coarsely tubercular . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Litoria inermis (p. 75) Skin of back slightly tubercular . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Litoria pallida (p. 80) 31

Body squat; legs short; dorsal glands conspicuous (Figure 6) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32 (Uperoleia) 148

KEY TO THE KIMBERLEY REGION

Body elongated; legs long; dorsal glands inconspicuous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41 (Crinia) 32

Upper jaw toothed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33 Upper jaw not toothed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35 33

Small but prominent papilla on heel. . . . . . . . . . . . . . . . . . . . . . . . . . Uperoleia mjobergii (p. 129) No papilla on heel. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34 34

Small body size (to 22 mm S-V). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Uperoleia micra (p. 126) Medium body size (~ 30 mm S-V) . . . . . . . . . . . . . . . . . . . . . . . . . . Uperoleia marmorata (p. 125) 35

Thigh and groin markings red . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36 Thigh and groin markings not red . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37 36

Back brown and coarsely tubercular . . . . . . . . . . . . . . . . . . . . . . . . . . . Uperoleia minima (p. 128) Back slatey-grey and densely covered with small, orange-tipped, conical tubercles. . . . . . . . . . . . . . . . . . . . . . Uperoleia trachyderma (p. 132) 37

Thigh and groin orange . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Uperoleia borealis (p. 121) Thigh and groin not orange . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38 38

Webbing between toes extensive, reaches about one quarter of the way up the fourth toe . . . . . . . . . . . . . . . . . . . . . Uperoleia talpa (p. 131) Webbing between toes, if present, confined to base. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39 39

Back coarsely tubercular . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Uperoleia aspera (p. 120) Back smooth or sparsely tubercular. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40 40

Call an explosive tick; glands golden and connect along sides; often a median stripe on head. . . . . . . . . . . . . . . . . . . . . Uperoleia lithomoda (p. 124) Call a slow rasp; no golden glands on sides; no median head stripe. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Uperoleia crassa (p. 122) 41

Network of bluish-white tubercles on back; males with wide flanges on fingers. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Crinia fimbriata (p. 32) No network of bluish-white tubercles on back; fingers without flanges . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 42 42

Squarish central blotch on back; call a rattle. . . . . . . . . . . . . . . . . . . . . . . . Crinia bilingua (p. 30) Broken blotches on back; call a chirp . . . . . . . . . . . . . . . . . . . . . . . . . . . . Crinia deserticola (p. 31) 149

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Nieden, F. (1923) Amphibia. Anura I. Subordo Aglossa und Phaneroglossa. Sectio 1 Arcifera. Das Tierreich 46: 1–584. Packer, W.C. (1963) Dehydration, hydration and burrowing behaviour in Heleioporus eyrei (Gray) (Leptodactylidae). Ecology 44: 643–651. Packer, W.C. (1966) Embryonic and larval development of Heleioporus eyrei (Amphibia: Leptodactylidae). Copeia 1966: 92–97. Parker, H.W. (1940) The Australasian frogs of the family Leptodactylidae. Novit. Zool. 42: 1–106. Peters, W. (1863) Übersicht der von Hrn. Richard Schomburgk an das zoologische Museum eingesandten Amphibien, aus Buchsfelde bei Adelaide in Südaustralien. Mber. K. Preuss. Akad. Wiss. Berl. 1863: 228–236. Peters, W. (1867) Herpetologische Notizen. Mber. K. Preuss. Akad. Wiss. Berl. 1867: 13–37. Philipp, G.A. (1958) Myobatrachus gouldii in the coastal hills near City Beach. West. Aust. Nat. 6: 131–132. Plötner, J., Kohler, K., Uzzell, T. and Beerli, P. (2007) Molecular systematics of amphibians. Pp. 2673–2756. In H. Heatwole: Amphibian Biology, Vol. 7. Surrey Beatty & Sons, Chipping Norton, NSW. Pramuk, J.B., Robertson, T., Sites, J.W., Jr. and Noonan, B.P. (2008) Around the world in 10 million years: biogeography of the nearly cosmopolitan true toads (Anura: Bufonidae). Global. Ecol. Biogeogr. 17: 72–83. Read, K., Keogh, J.S., Scott, I.A.W., Roberts, J.D. and Doughty, P. (2001) Molecular phylogeny of the Australian frog genera Crinia and Geocrinia and allied taxa (Anura:Myobatrachidae). Mol. Phyl. Evol. 21: 294–308. Read, J. (1999a) Diet and causes of mortality of the trilling frog (Neobatrachus centralis). Herpetofauna 29: 13–18. Read, J. (1999b) Abundance and recruitment patterns of the trilling frog (Neobatrachus centralis) in the Australian arid zone. Aust. J. Zool 47: 393–404. Roberts, J.D. (1981) Terrestrial breeding in the Australian leptodactylid frog Myobatrachus gouldii (Gray). Aust. Wildl. Res. 8: 451–462. Roberts, J.D. (1984) Terrestrial egg deposition and direct development in Arenophryne rotunda Tyler, a myobatrachid frog from coastal sand dunes at Shark Bay, W.A. Aust. Wildl. Res. 11: 191–200. Roberts, J.D. (1985) Population estimates for Arenophryne rotunda: is the round frog rare? In G. Grigg, R. Shine and H. Ehmann (Eds). Biology of Australasian frogs and reptiles. pp. 463–467. (Royal Zoological Society of New South Wales, Sydney). Roberts, J.D. (1990) The biology of Arenophryne rotunda (Anura: Myobatrachidae): a burrowing frog from Shark Bay, Western Australia. In P.F. Beery, S.D. Bradshaw and B.R. Wilson (Eds). ‘Research in Shark Bay. Report of the France-Australe Bicentenary Expedition Committee. pp. 287–297. (Western Australian Museum, Perth). Roberts, J.D. (1997a) Call evolution in Neobatrachus (Anura: Myobatrachidae). Speculation on tetraploid origins. Copeia (1997): 791–801. Roberts, J.D. (1997b) Geographic variation in calls of males and determination of species boundaries in tetraploid frogs of the Australian genus Neobatrachus (Myobatrachidae). Aust. J. Zool. 45: 95–112. Roberts, J.D., Horwitz, P., Wardell-Johnson, G., Maxson, L.R. and Mahony, M.J. (1997) Taxonomy, relationships and conservation of a new genus and species of myobatrachid frog from the high rainfall region of southwestern Australia. Copeia 1997: 373–381. 153

FROGS OF WESTERN AUSTRALIA Roberts, J.D., Mahony, M., Kendrick, P. and Majors, C.M. (1991) A new species of burrowing frog, Neobatrachus (Anura: Myobatrachidae), from the eastern wheatbelt of Western Australia. Rec. West. Aust. Mus. 15: 23–32. Roberts, J.D. and Majors, C.M. (1993) Range extensions, range definitions and call structures for frogs from Western Australia. Rec. West. Aust. Mus. 16: 315–322. Roberts, J.D. and Maxson, L.R. (1986) Phylogenetic relationships in the genus Limnodynastes (Anura: Myobatrachidae). Aust J. Zool. 34: 561–573. Roberts, J. D. and Maxson, L. R. (1989) A molecular perspective on relationships of Australian Pseudophryne (Anura: Myobatrachidae). Syst. Zool. 38: 154–165. Roberts, J.D., Standish, R., Byrne, P. and Doughty, P. (1999) Synchronous polyandry and multiple paternity in the frog Crinia georgiana (Anura: Myobatrachidae). Anim. Behav. 57: 721–726. Roberts, J.D. and Wardell-Johnson, G. (1995) Call differences between peripheral isolates of the Geocrinia rosea complex (Anura: Myobatrachidae) in southwestern Australia. Copeia 1995: 899–906. Roberts, J.D., Wardell-Johnson, G. and Barendse, W. (1990) Extended descriptions of Geocrinia vitellina and Geocrinia alba (Anura: Myobatrachidae) from Southwestern Australia, with comments on the status of G. lutea. Rec. West. Aust. Mus. 14: 427–437. Roberts, J.D. and Watson, G.F. (1993) Biogeography and phylogeny of the Anura. In C. J. Glasby, G. J. B. Ross and P. L. Beesley (Eds). Fauna of Australia. Vol. 2A Amphibia and Reptilia. Pp. 35–40. (Australian Government Publishing Service, Canberra). Schäuble, C.S., Moritz, C. and Slade, R.W. (2000) A molecular phylogeny for the frog genus Limnodynastes (Anura: Myobatrachidae). Mol. Phylo. Evol. 16: 379–391. Schlegel, H. (1850) Description of a new genus of batrachians from the Swan River. Proc. Zool. Soc., Lond. 1850: 9–10. Seymour, R.S. and Roberts, J.D. (1995) Oxygen uptake by the aquatic eggs of the Australian frog Crinia georgiana. Physiol. Zool. 68: 206–222. Seymour, R.S., Roberts, J.D., Mitchell, N.J. and Blaylock, A.J. (2001) Influence of environmental oxygen on development and hatching of aquatic eggs of the Australian frog, Crinia georgiana. Physiol. Biochem. Zool. 73: 501–507. Shea, G.M. and Johnston, G.R. (1987) A new species of Notaden (Anura: Leptodactylidae) from the Kimberley Division of Western Australia. Trans. R. Soc. S. Aust. 112: 29–37. Slater, P. and Main, A.R. (1963) Notes on the biology of Notaden nichollsi Parker (Anura, Leptodactylidae). West. Aust. Nat. 8: 163–166. Smith, M. and Roberts, J.D. (2003a) Call structure may affect mating success in the quacking frog Crinia georgiana (Anura: Myobatrachidae). Behav. Ecol. Sociobiol. 53: 221–226. Smith, M. and Roberts, J.D. (2003b) An experimental examination of female preference patterns for components of the male advertisement call in the quacking frog (Crinia georgiana). Behav. Ecol. Sociobiol. 53: 221–226. Smith, M. and Roberts, J.D. (2003c) No sexual dimorphism in the frog Crinia georgiana (Anura: Myobatrachidae): an examination of pre and post–maturational growth. J. Herpetol. 37: 132–137. Smith, M. and Roberts, J.D. (2003d) Call repertoire of an Australian treefrog, Litoria adelaidensis (Anura, Hylidae). J. Roy. Soc. West. Aust. 86: 91–95. 154

Smith, M.J., Roberts, J.D., Hammond, T.J. and Davis, R.A. (2003) Intra-specific variation in the advertisement call and body size relationships in mated pairs of the sunset frog Spicospina flammocaerulea (Anura: Myobatrachidae). J. Herp. 37: 285–291. Smith, M.J., Withers, P.C. and Roberts, J.D. (2003a) Reproductive energetics and behavior of an Australian myobatrachid frog Crinia georgiana. Copeia 2003: 248– 254. Spencer, B. (1896) Report on the works of the Horn Scientific Expedition to Central Australia I. Introduction, Narrative, Summary of results, supplement to zoological report. Amphibia (Melville, Muller and Slade: Melbourne). Steindachner, F. (1867) Amphibien. In Reise der osterreichischen Fregatte Novara um die Erde in den Jahren 1857, 1858, 1859. Zoologie 1: 1–70. Straughan, I.R. (1969) Hyla inermis (Peters) a species hitherto erroneously referred to the leptodactylid genus Cyclorana (Anura: Hylidae: Leptodactylidae). Zool. Meded., Leiden 43: 207–212 Thompson, G.G., Withers, P.C., Mcmaster, K.A. and Cartledge, V.A. (2005) Burrows of desert-adapted frogs, Neobatrachus aquilonius and Notaden nichollsi. J. Roy. Soc. W. Aust. 88:17–23. Tschudi, J.J. von. (1838) Classification der Batrachier mit Berücksichtigung der fossilen Thiere dieser Abtheilung der Reptilien. (Petitpierre, Neuchâtel, Germany). Tyler, M.J. (1968) A taxonomic study of hylid frogs of the Hyla lesueuri complex occurring in north-western Australia. Rec. S. Aust. Mus. 15: 711–272. Tyler, M.J. (1969) A synopsis of the frogs of the genus Hyla of northwestern Australia, with the description of a new species. Rec. S. Aust. Mus. 16: 1–11. Tyler, M.J. (1972) Superficial mandibular musculature, vocal sacs and the phylogeny of Australo-Papuan leptodactylid frogs. Rec. S. Aust. Mus. 16: 1–20. Tyler, M.J. (1976) A new genus and two new species of leptodactylid frogs from Western Australia. Rec. W. Aust. Mus. 4: 45–52. Tyler, MJ. (1978) Amphibians of South Australia. (Government Printer: Adelaide). Tyler, M.J. (1979) Herpetofaunal relationships of South America with Australia. In W. E. Duellman (Ed.) The South American Herpetofauna: its origin, evolution and dispersal. Mus. Nat. Hist. Univ. Kansas Monogr. 7: 73– 106. Tyler, M.J. (1983) Neobatrachus sutor Main: a frog new to the fauna of South Australia. Trans. Roy. Soc. S. Aust. 107: 129–130. Tyler, M.J. (1985) Quaternary fossil frogs from Skull Cave and Devil’s Lair in the extreme southwest of Western Australia. Rec. W. Aust. Mus. 12: 233–240. Tyler, M.J. (1990) Geographic distribution of the fossorial hylid frog Cyclorana platycephala (Günther) and the taxonomic status of C. slevini Loveridge. Trans. R. Soc. S. Aust. 114: 81–85. Tyler, M.J. (1994) Australian frogs. A natural history. (Reed: Melbourne). Tyler, M.J. (1996) Frogs as pets: a guide to keeping the Australian Green Tree Frog (Litoria caerulea). (Graphic Print Group, Adelaide). Tyler, M.J. and Crook, G.A. (1987) Technical memorandum No. 19. Supervising Scientist for the Alligator Rivers Region, Australian Government Publishing Service, Bondi Junction, NSW.

REFERENCES Tyler, M.J., Crook, G.A. and Davies, M. (1983a) Reproductive biology of the frogs of the Magela Creek System, Northern Territory. Rec. S. Aust. Mus. 18: 415–440. Tyler, M.J. and Davies, M. (1978) Species-groups within the Australopapuan hylid frog genus Litoria Tschudi. Aust. J. Zool. Suppl. Ser. No. 63: 1–47. Tyler, M.J. and Davies, M. (1979a) Foam nest construction by Australian Leptodactylid frogs (Amphibia, Anura, Leptodactylidae). J. Herp. 13: 509–510. Tyler, M.J., and Davies, M. (1979b) A new species of cavedwelling, hylid frog from Mitchell Plateau, Western Australia. Trans. Roy. Soc. S. Aust. 103: 149–153. Tyler, M.J. and Davies M. (1980) Systematic status of Kankanophryne Heyer and Liem (Anura: Leptodactylidae). Trans. R. Soc. S. Aust. 104: 17–20. Tyler, M.J. and Davies, M. (1993) Family Hylidae. In In C.J. Glasby, G.J.B. Ross and P.L. Beesley (Eds). Fauna of Australia. Vol. 2A Amphibia and Reptilia. Pp. 58–63. (Australian Government Publishing Service, Canberra). Tyler, M.J. and Davies, M. (2000) Developmental biology and larval morphology of the frog Limnodynastes depressus Tyler (Myobatrachidae: Limnodynastinae). Trans. Roy. Soc. S. Aust. 124: 169–175. Tyler, M.J., Davies, M. and King, M. (1978) The Australian frog Chiroleptes dahlii Boulenger: Its systematic position, morphology, chromosomes and distribution. Trans. R. Soc. S. Aust. 102: 17–23. Tyler, M.J. Davies, M. and Martin, A.A. (1977) A new species of large green tree frog from northern Western Australia. Trans. R. Soc. S. Aust. 101: 133–138. Tyler, M.J., Davies, M. and Martin, A.A. (1981a) Australian leptodactylid frogs of the genus Uperoleia. Aust. J. Zool. Suppl. Ser. 79: 1–64. Tyler, M.J., Davies, M. and Martin, A.A. (1981b) New and rediscovered species of frogs from the Derby-Broome area of Western Australia. Rec. West. Aust. Mus. 9: 147–172. Tyler, M.J., Davies, M. and Martin, A.A. (1981c) Frog fauna of the Northern Territory: new distributional records and the description of a new species. Trans. R. Soc. S. Aust. 105:149–154. Tyler, M.J., Davies, M. and Martin, A.A. (1982) Biology, morphology and distribution of the Australian fossorial frog Cyclorana cryptotis (Anura: Hylidae). Copeia 1982(3): 260–264. Tyler, M.J., Davies, M. and Martin A.A. (1983b) The frog fauna of the Barkly Tableland, Northern Territory. Trans. R. Soc. S. Aust. 107: 237–242. Tyler, M.J. and Ledo, W.W. (1973) The type locality of the leptodactylid frog Neobatrachus centralis Parker. S. Aust. Nat. 47: 75–76. Tyler, M.J. and Martin, A.A. (1975) Australian leptodactylid frogs of the Cyclorana australis complex. Trans. R. Soc. S. Aust. 99: 93–99. Tyler, M.J. and Martin, A.A. (1977) Taxonomic studies of some Australian leptodactylid frogs of the genus Cyclorana Steindachner. Rec. S. Aust. Mus. 17: 261–276. Tyler, M.J., Martin, A.A. and Davies, M. (1979) Biology and systematics of a new limnodynastine genus (Anura: Leptodactylidae) from north-western Australia. Aust. J. Zool. 27: 135–150. Tyler, M.J., Roberts, J.D. and Davies, M. (1980) Field observations on Arenophryne rotunda Tyler, a leptodactylid frog of coastal sandhills. Aust. Wildl. Res. 7: 295–304. Van Beurden, E. (1979) Gamete development in relation to season, moisture, energy reserve, and size in the Australian water-holding frog Cyclorana

platycephalus. Herpetologica 35: 370–374. Van Beurden, E. (1982) Desert adaptations of Cyclorana platycephala: a holistic approach to desert-adaptation in frogs. In W. R. Barker and P. J. M. Greenslade (Eds.). Evolution of the flora and fauna of arid Australia. pp. 235–240. (Peacock Publications, Adelaide, SA). Wabnitz, P.A., Walters, H., Tyler, M.J., Wallace, J.C. and Bowie, J.H. (1998) First record of host defence peptides in tadpoles. The Magnificent Tree Frog Litoria splendida. J. Peptide Res. 52: 477–481. Wagler, J.G. (1828) Vorläufige Uebersicht des Geruftes, sowie Untungigung feines Systema amphibiorum. Isis von Oken 21: 859–861. Warburg, M.R. (1967) On thermal and water balance of three Central Australian frogs. Comp. Biochem. Physiol. 20: 27–43. Wardell-Johnson, G. and Roberts, D. (1989) Endangered! Forest frogs. Landscope 5: 17. Wardell-Johnson, G. and Roberts, J.D. (1993) Biogeographic barriers in a subdued landscape: the distribution of the Geocrinia rosea (Anura: Myobatrachidae) complex in south-western Australia. J. Biogeog. 20: 95–108. Wardell-Johnson, G. and Roberts, J.D. (1996) Morphological variation in the Geocrinia rosea (Anura: Myobatrachidae) complex in south-western Australia. In S. Hopper, J. Chappil, M. Harvey and A. George (Eds.). Gondwanan Heritage: Past, Present and Future of the Western Australian Biota. pp. 172–178. (Surrey-Beatty, ChippingNorton, NSW). Watson, G.F. and Gerhardt, H.C. (1997) The breeding biology and advertisement call of Litoria splendida Tyler, Davies & Martin. Trans. R. Soc. S. Aust. 121: 119–124. Watson, G.F. and Martin, A.A. (1973) Life history, larval morphology and relationships of Australian leptodactylid frogs. Trans. R. Soc. S. Aust. 97: 33–45. Watson, J.A.L. and Saunders, L.M. (1959) Observations on the reproductive system of the female of Myobatrachus gouldii (Gray). W. Aust. Nat. 7: 1–6. Werner, F. (1914) Amphibia. In: Michaelsen, W., and Hartmeyer, R. (Eds.). Die Fauna Sudwest-Australiens. 4: 403–426. White, J. (1790) Journal of a Voyage to New South Wales, with sixty five plates of non-descript animals, birds, lizards, serpents, curious cones of trees and other natural productions. (Debrett, London, UK). Withers, P.C. (1993) Metabolic depression during aestivation in the Australian frogs, Neobatrachus and Cyclorana. Aust. J. Zool. 41: 467–473. Withers, P.C. (1995a) Cocoon formation and structure in aestivating Australian desert frogs, Neobatrachus and Cyclorana. Aust. J. Zool. 43: 429–441. Withers, P.C. (1995b) Evaporative water loss and colour change in the Australian desert tree frog Litoria rubella (Amphibia: Hylidae). Rec. West. Aust. Mus. 17: 277– 281. Withers, P. C. (1998) Evaporative water loss and the role of cocoon formation in Australian frogs. Aust. J. Zool. 46: 405–418 Withers, P.C. and Roberts, J.D. (1993) Water balance response in the terrestrial Western Australian frog Myobatrachus gouldii (Anura: Myobatrachidae). J. Roy. Soc. W. Aust. 76: 49–52. Withers, P.C. and Thompson, G.G. (2000) Cocoon formation and metabolic depression by the aestivating hylid frogs Cyclorana australis and Cyclorana cultripes (Amphibia: Hylidae). J. Roy. Soc. West. Aust. 83: 39–40. 155

ACKNOWLEDGEMENTS We thank L. A. Smith and R. E. Johnstone for contributions to earlier editions of this field guide; their knowledge and spirit of discovery were essential for this series. Fieldwork in the Kimberley region was made possible by grants from the Australian Research Council, Utah Foundation and Alcoa of Australia as part of the Western Australian Museum’s Alcoa Frog Watch program. Permits for fieldwork, and for the transfer of specimens to South Australia for further study, were provided by the Department of Environment and Conservation (formerly Conservation and Land Management). We are grateful to the following colleagues for help in the field, laboratory and in discussions: M. Davies, A. Martin, G. Watson, D. Williams, C. Miller, J. Bishop, D. Roberts, B. Maryan, M. Anstis, D. Pearson, J. Francis, C. Mills, S. Donnellan, L. Price, P. Oliver, D. Moore, S. Keogh, D. Edwards, M. Dziminski, C. Stevenson, P. Kendrick, M. Barrett, R. Barrett, V. Kessner, N. Mitchell, J. Prefumo, M. Peterson and G. Harold. We are also indebted to the following who helped our investigations in various ways: G. Storr, D. King, J. Caratti, K. Mahoney, M. Mahony, E. Lightburn, G. Mewett, B. Pennington, J. Dell, K. Aplin, M. Cowan, R. How, M. Greenlees, B. Phillips, P. Horner, D. Trembath, P. Couper, A. Amey and J. Rolfe. The excellent photographs of frogs were generously provided by B. Maryan, G. Harold, J. Francis, M. Peterson, G. Gaikhorst, M. Anstis, B.B. Wells (DEC), M. Cowan, D. Moore, C. McCarthy (BMNH), H. Cogger, L. Price and S. Donnellan. All copyright for images is with the photographers or the Western Australian Museum. We thank L. Trueb and N. Guthrie for the illustrations and C. Stevenson for preparing the distribution maps from specimens in the collections of the Western Australian Museum. We thank J. D. Roberts who read through an entire draft of the manuscript and G. Shea for numerous taxonomic recommendations, including extensive comments on etymology.

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Michael J. Tyler has studied frogs for 50 years and published the results throughout the world. He is Associate Professor of Ecology and Evolutionary Biology and Visiting Research Fellow at the University of Adelaide. In 1997 he won the Michael Daley Eureka Prize for the Promotion of Science. Paul Doughty is Curator in Herpetology at the Western Australian Museum and coordinator of the Alcoa Frog Watch program, and he is an active advocate for frog education, through public talks and radio and newspaper interviews.

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INDEX Unlike Latin or Greek-based scientific names, common names of species are not fixed by a strict code of rules. As such, they can vary widely across areas and regions. In keeping with recent attempts to provide common names for all species for the public (e.g. Ingram et al. 1983; CSESN 2008), we have updated the list of common names in this field guide in an attempt to stabilize common name useage. We have usually not kept names that merely repeat scientific names coined after people. Many of these names persist because the species are not well-known so that over time they do not take on a common name that reflects the species’ appearance, call, region of occurrence or other relevant features (most scientific names relate to the species’ biology in some way). For example, Metacrinia nichollsi has been changed from Nicholls’ Toadlet to the Forest Toadlet. We feel the latter name conveys something about the biology of the species to people outside the field of herpetology, and is therefore a more useful common name. Aquatic Frog..................................... 74 Arenophryne rotunda.......................... 26 Arenophryne xiphorhyncha.................. 27 Bilingual Froglet................................ 30 Black-shinned Rocket Frog................ 85 Blacksoil Toadlet............................. 132 Bleating Froglet................................. 37 Bufo marinus................................... 135 Bumpy Rocket Frog.......................... 75 Cane Toad....................................... 135 Carpenter Frog...................................65 Cave Frog.......................................... 71 Centralian Burrowing Frog.............. 109 Chattering Rock Frog........................ 84 Chirping Froglet................................ 31 Clicking Froglet................................ 35 Crawling Toadlet............................. 113 Crinia bilingua.................................. 30 Crinia deserticola................................ 31 Crinia fimbriata................................. 32 Crinia georgiana................................. 33 Crinia glauerti................................... 35 Crinia insignifera............................... 36 Crinia pseudinsignifera....................... 37 Crinia subinsignifera.......................... 38 Cyclorana australis.............................. 40 Cyclorana cryptotis.............................. 41 Cyclorana cultripes.............................. 42 Cyclorana longipes.............................. 43 Cyclorana maini................................. 44 Cyclorana platycephala........................ 45 Cyclorana vagitus................................ 46 Derby Toadlet................................. 120 Desert Spadefoot............................. 105 Desert Trilling Frog........................... 96 Fat Toadlet...................................... 122 Flat-headed Frog............................... 63 Forest Toadlet.................................... 89 Geocrinia alba.................................... 48 Geocrinia leai..................................... 49 Geocrinia lutea................................... 50 Geocrinia rosea................................... 51 Geocrinia vitellina.............................. 52 Giant Frog......................................... 40 Glandular Toadlet........................... 123 Gorge Toadlet.................................. 112 Green Tree Frog................................. 70 Heleioporus albopunctatus................... 54 Heleioporus barycragus........................ 55 Heleioporus eyrei................................. 56 Heleioporus inornatus......................... 57 Heleioporus psammophilus.................. 58 Hidden-ear Frog................................ 41 Hooting Frog.................................... 55 Humming Frog................................. 99 Javelin Frog....................................... 77 Kimberley Froglet.............................. 32 158

Kimberley Spadefoot....................... 106 Knife-footed Frog.............................. 42 Limnodynastes convexiusculus.............. 62 Limnodynastes depressus...................... 63 Limnodynastes dorsalis........................ 64 Limnodynastes lignarius...................... 65 Litoria adelaidensis............................. 68 Litoria bicolor.................................... 69 Litoria caerulea.................................. 70 Litoria cavernicola.............................. 71 Litoria coplandi.................................. 72 Litoria cyclorhyncha............................ 73 Litoria dahlii..................................... 74 Litoria inermis................................... 75 Litoria meiriana................................. 76 Litoria microbelos............................... 77 Litoria moorei.................................... 78 Litoria nasuta..................................... 79 Litoria pallida.................................... 80 Litoria rothii...................................... 81 Litoria rubella.................................... 82 Litoria splendida................................ 83 Litoria staccato................................... 84 Litoria tornieri................................... 85 Litoria watjulumensis......................... 86 Little Red Tree Frog........................... 82 Long-footed Frog.............................. 43 Magnificent Tree Frog....................... 83 Main’s Frog....................................... 44 Marbled Frog.................................... 62 Marbled Toadlet.............................. 125 Metacrinia nichollsi............................ 89 Moaning Frog................................... 56 Mole Toadlet................................... 131 Motorbike Frog................................. 78 Myobatrachus gouldii.......................... 91 Neobatrachus albipes........................... 94 Neobatrachus aquilonius..................... 95 Neobatrachus centralis......................... 96 Neobatrachus fulvus............................ 97 Neobatrachus kunapalari.................... 98 Neobatrachus pelobatoides................... 99 Neobatrachus sutor........................... 100 Neobatrachus wilsmorei.................... 101 Northern Burrowing Frog................. 95 Northern Dwarf Tree Frog................. 69 Northern Laughing Tree Frog............ 81 Northern Sandhill Frog..................... 26 Northern Spadefoot........................ 104 Northern Toadlet............................. 121 Northwest Toadlet........................... 130 Notaden melanoscaphus.................... 104 Notaden nichollsi.............................. 105 Notaden weigeli................................ 106 Orange-bellied Frog.......................... 52 Ornate Burrowing Frog................... 108 Pale Rocket Frog............................... 80

Platyplectrum ornatum..................... 108 Platyplectrum spenceri....................... 109 Plonking Frog................................. 101 Pseudophryne douglasi....................... 112 Pseudophryne guentheri..................... 113 Pseudophryne occidentalis.................. 114 Quacking Frog.................................. 33 Rattling Froglet................................. 35 Rock Frog......................................... 72 Rockhole Frog................................... 76 Roseate Frog...................................... 51 Sand Frog.......................................... 58 Sheep Frog........................................ 44 Shoemaker Frog.............................. 100 Slender Tree Frog............................... 68 Small Toadlet .................................. 128 South Coast Froglet........................... 38 Southern Sandhill Frog...................... 27 Spicospina flammocaerulea................ 117 Splendid Tree Frog............................ 83 Spotted-thighed Frog........................ 73 Squelching Froglet............................. 36 Stonemason Toadlet........................ 124 Striped Rocket Frog........................... 79 Sunset Frog..................................... 117 Tanami Toadlet............................... 127 Tawny Trilling Frog........................... 97 Ticking Frog...................................... 49 Tiny Toadlet.................................... 126 Turtle Frog........................................ 91 Uperoleia aspera............................... 120 Uperoleia borealis............................. 121 Uperoleia crassa................................ 122 Uperoleia glandulosa......................... 123 Uperoleia lithomoda......................... 124 Uperoleia marmorata........................ 125 Uperoleia micra................................ 126 Uperoleia micromeles........................ 127 Uperoleia minima............................ 128 Uperoleia mjobergii.......................... 129 Uperoleia russelli.............................. 130 Uperoleia talpa................................. 131 Uperoleia trachyderma...................... 132 Wailing Frog..................................... 46 Walpole Frog..................................... 50 Water-holding Frog........................... 45 Western Banjo Frog........................... 64 Western Spotted Frog........................ 54 Western Toadlet.............................. 114 West Kimberley Toadlet.................. 129 Wheatbelt Frog................................. 98 White-bellied Frog............................ 48 White-footed Trilling Frog................ 94 Whooping Frog................................. 57 Woodworker Frog..............................65 Wotjulum Frog...................................86 Yellow-bellied Frog............................ 52

Published by the Western Australian Museum 49 Kew Street, Welshpool, Western Australia 6106 www.museum.wa.gov.au First published 1984 Second edition 1994 Third edition 2000 Fourth edition 2009 Digital edition released 2013. Copyright © Western Australian Museum, 2009 Designed by Cathie Glassby and Michael Neiman Printed by South Wind Productions, Singapore The National Library of Australia Cataloguing-in-Publication entry: Tyler, Michael J., 1937– Frogs of Western Australia 4th edition. ISBN 978 1 920843 40 3 (pbk.) ISBN 978 1 920843 91 5 (epdf) I. Frogs – Western Australia. II. Doughty, P. (Paul), 1965–, III. Western Australian Museum. IV. Title. 597.809941

Front Cover: Litoria moorei (© Jiri Lochman, Lochman Transparencies) Back Cover (from the top): Platyplectrum ornatum (G. Harold); Litoria microbelos (G. Harold); Litoria moorei (G. Harold).

Western Australia is home to more than 80 of the known frog species in Australia and new frogs are still being discovered. This comprehensive guide will appeal to general and specialist readers and to students; to anyone with an interest in Australia’s frogs, which are an important barometer of environmental health. It presents up-to-date information on all species, including their distribution and habitat preferences, and features extensive colour photographs and location maps to assist with identification.