Status of Conservation and Decline of Amphibians: Australia, New Zealand, and Pacific Islands

Amphibians are among the most threatened groups of animals on earth. In part due to their highly permeable skin, amphibians are highly sensitive to environmental changes and pollution and provide an early-warning system of deteriorating environmental conditions. The more we learn about the impact of environmental changes on amphibians, the better we as humans will be able to arrest their demise, and our own.

Status of Conservation and Decline of Amphibians brings together the current knowledge on the status of the unique frogs of Australia, New Zealand, and the Pacific. Although geographically proximate, each region presents unique challenges and opportunities in amphibian research and conservation. This book contributes to an understanding of the current conservation status of the amphibians of each region, aims to stimulate research into halting amphibian declines, and provides a better foundation for making conservation decisions. It is an invaluable reference for environmental and governmental agencies, researchers, policy-makers involved with biodiversity conservation, and the interested public.

• Language: English
• Publisher: CSIRO PUBLISHING
• Release date: Jun 1, 2018
• ISBN: 9781486308408

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STATUS OF CONSERVATION AND DECLINE OF AMPHIBIANS

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Front cover: (top) Neobatrachus sutor, Stephen Mahony; (bottom) mountain ash forest, David Blair; (thumbnails, left to right) Cornufer vogti, Robert Fisher, U.S. Geological Survey; Notaden nicholsii, Stephen Mahony; Leiopelma archeyi, Phil Bishop

Back cover: (left to right) Cyclorana platycephela, Stephen Mahony; Litoria cyclorhyncha, JD Roberts; Myobatrachus gouldii, Stephen Mahony

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STATUS OF CONSERVATION AND DECLINE OF

AMPHIBIANS

AUSTRALIA, NEW ZEALAND, AND PACIFIC ISLANDS

Editors: Harold Heatwole and Jodi J. L. Rowley

Dedication

The name of Dr Harold (Hal) Cogger has been synonymous with Australian herpetology for several generations. His monumental treatise Reptiles & Amphibians of Australia won the coveted Whitley Award and has been the mainstay of professionals, and an inspiration for amateur herpetologists and naturalists. Not only is it a classic work – seldom, if ever, equalled for any other geographic region – but he has revised it frequently so that it has served as a continuous, authoritative, publicly available source of the latest information on the frogs, lizards, snakes, turtles, and crocodiles of Australia. The first edition in 1975 listed the 664 species then known for the country. The discovery of new species, at least five described by Dr Cogger himself, has burgeoned over the decades since then, with seven successive editions sequentially yielding additional species, until the latest one in 2014 was double in size and peaked at 1218 species! Without this manual, it would have been nearly impossible for even the professional herpetologist to keep abreast of so many advances in knowledge, or to be able to readily identify species encountered in the field. Dr Cogger is an excellent photographer and he took all but a few of the more than 840 coloured pictures gracing his guide. This book alone constitutes a lasting legacy of enormous proportions, but it is merely one of his many achievements.

In collaboration with his assistant, Elizabeth Cameron, and his wife Heather Cogger, he (1983) produced a catalogue Amphibia and Reptilia as the first volume of the Zoological Catalogue of Australia. In addition to these books, he has published about 160 professional scientific articles. His thorough knowledge of the Australian herpetofauna is evident in his treatment of amphibians in Chapter 2 of the present volume. He has had one genus (Coggeria), seven species (Coggeria naufragus, Ctenotus coggeri, Lampropholis coggeri, Oedura coggeri, Aprasia haroldi, Delma haroldi, Lerista haroldi, and one subspecies, Amphibolurus nobbi coggeri of lizard, as well as one species of sea snake, Hydophis coggeri, named in his honour.

Harold Cogger is a consummate field biologist with wide experience in New Guinea and Australia and its islands (Cape York, Swain Reefs, Central Australia, Arnhem Land, Cocos-Keeling, Torres Straits, Ashmore Reef). He has participated in various international field expeditions: for example, to Australasia to study sea snakes aboard the Alpha Helix, a vessel of Scripps Institution of Oceanography, on two expeditions, as well as several expeditions sponsored by the Japanese Ministry of Science, Education, and Culture to study sea snakes in the western and south-western Pacific. These expeditions were memorable for the comradery among the participants from Japan, France, Australia, and the United States.

Hal Cogger started out as a cadet preparator at the Australian Museum in 1952 and rose through the ranks of that institution progressively as Assistant Curator (birds, reptiles, and amphibians), Curator (reptiles and amphibians), Senior Research Scientist, and Deputy Director. He retired in 1995 but has remained active in research as a John Evans Memorial Fellow of the Australian Museum up to the present and was Conjoint Professor in Sustainable Resource Management at the University of Newcastle from 1997 until 2002. Many of his later contributions have been in the field of conservation as: Chairman of the Australasian Reptile and Amphibians Specialist Group, Survival Commission, International Union of the Conservation of Nature (1991–2000); Chairman of the Australian Biological Resources Study Advisory Committee (1992–1997); member of the Endangered Fauna (Interim Protection) Act 1991 Scientific Committee (1992–1995); and the Australian Biological Diversity Advisory Committee (1991–1993). He was the lead researcher and the author of the Action Plan for Australian Reptiles in 1993.

In modern times, scientists often are competitive, but Harold Cogger treats science as a cooperative venture and generously and freely shares his knowledge, expertise, and enthusiasm about reptiles and amphibians. His immersion in his chosen field is perhaps best measured by his expression on one occasion that ‘You haven’t really lived until you’ve heard a full-throated chorus of frogs’. A glimpse of what stimulated Hal Cogger to embark on such a successful career and develop exuberance for the hobby he turned into a profession is captured by a published interview with him by Neville Burns in 2014.

I have kept secret the fact that I planned to honour Hal by dedicating this book to him. The first he will be aware of it is when he opens his personal copy and stares at his own face peering from the page!

Harold Heatwole

1 August 2017

Contents

This book is part of the Amphibian Biology series, Volume 11, Status of Conservation and Decline of Amphibians: Eastern Hemisphere. The chapters in this book (Chapters 1–14) are consecutive chapters in Part 6 (Australia, New Zealand, and Pacific Islands); numbers of chapters in parentheses (68–81) are the same chapters numbered consecutively across all parts of Volume 11.

Dedication

Contents of Previous Parts of Volume 11

Contributors to Part 6

Preface

Chapter 1 (68). Introduction

Harold Heatwole and Jodi J. L. Rowley

Chapter 2 (69). A Brief Demographic Overview of Australia’s Native Amphibians

Harold G. Cogger

Chapter 3 (70). Status of Decline and Conservation of Frogs in the Wet Tropics of Australia

Ross A. Alford and Jodi J. L. Rowley

Chapter 4 (71). Frogs of the Monsoon Tropical Savannah Regions of Northern Australia

Graeme R. Gillespie and J. Dale Roberts

Chapter 5 (72). An Update on Frog Declines from the Forests of Subtropical Eastern Australia

David Newell

Chapter 6 (73). Frog Declines and Associated Management Response in South-eastern Mainland Australia and Tasmania

David Hunter, Nick Clemann, David Coote, Graeme R. Gillespie, Greg Hollis, Ben Scheele, Annie Philips, and Matt West

Chapter 7 (74). The Status of Decline and Conservation of Frogs in Temperate Coastal South-eastern Australia

Frank Lemckert and Michael Mahony

Chapter 8 (75). Conservation of Frogs in South-western Australia

J. Dale Roberts

Chapter 9 (76). The Status of Decline and Conservation of Frogs in the Arid and Semi-arid Zones of Australia

Joanne Ocock and Skye Wassens

Chapter 10 (77). The Impact of an Invasive Amphibian: The Cane Toad Rhinella marina

Richard Shine

Chapter 11 (78). The Role of Ex-Situ Amphibian Conservation in Australia

Michael S. McFadden, Deon Gilbert, Kay Bradfield, Murray Evans, Gerry Marantelli, and Philip Byrne

Chapter 12 (79). Conservation of Frogs in Australia: State and Federal Laws

J. Dale Roberts

Plates

Chapter 13 (80). Status of Decline and Conservation of Frogs in New Zealand

Ben D. Bell and Phillip J. Bishop

Chapter 14 (81). Amphibians of the Pacific: Natural History and Conservation

George R. Zug and Robert N. Fisher

Index

Contents of previous parts of Volume 11, Amphibian Biology: Eastern Hemisphere

PART 1. CONSERVATION BIOLOGY OF AMPHIBIANS OF ASIA (EDITED BY HAROLD HEATWOLE AND INDRANEIL DAS), 2014. NATURAL HISTORY PUBLICATIONS (BORNEO), KOTA KINABALU. HARDBACK.

Chapter 1. Changes in Amphibian Populations in the Commonwealth of Independent States (Former Soviet Union)

Sergius L. Kuzmin and C. Kenneth Dodd Jr.

Chapter 2. Status of Conservation and Decline of Amphibians of Mongolia

Sergius L. Kuzmin

Chapter 3. Diversity and Conservation Status of Chinese Amphibians

Jianping Jiang, Feng Xie, and Cheng Li

Chapter 4. The Conservation of Amphibians in Korea

Daesik Park, Mi-Sook Min, Kelly C. Lasater, Jae-Young Song, Jae-Hwa Suh, Sang-Ho Son, and Robert H. Kaplan

Chapter 5. Conservation Status of Japanese Amphibians

Masafumi Matsui

Chapter 6. Status and Decline of Amphibians of Afghanistan

Indraneil Das

Chapter 7. Amphibians of Pakistan and their Conservation Status

Muhammad Sharif Khan

Chapter 8. Status and Decline of Amphibians of India

Indraneil Das and Sushil K. Dutta

Chapter 9. Sri Lankan Amphibians: Extinctions and Endangerment

Rohan Pethiyagoda, Kelum Manamendra-Arachchi, and Madhava Meergaskumbura

Chapter 10. Amphibians of the Maldives Archipelago

Indraneil Das

Chapter 11. Status, Distribution, and Conservation Issues of the Amphibians of Nepal

Karan B. Shah

Chapter 12. Status of Amphibian Studies and Conservation in Bhutan

Indraneil Das

Chapter 13. Status, Distribution and Conservation of the Amphibians of Bangladesh

A. H. M. Ali Reza

Chapter 14. Amphibian Conservation: Myanmar

Guinevere O. U. Wogan

Chapter 15. Decline of Amphibians in Thailand

Yodchaiy Chuaynkern and Prateep Duengkae

Chapter 16. Amphibian Conservation in Vietnam, Laos, and Cambodia

Jodi J. L. Rowley and Bryan L. Stuart

Chapter 17. Conservation Status of the Amphibians of Malaysia and Singapore

Indraneil Das, Norsham Yaakob, Jeet Sukumaran, and Tzi Ming Leong

Chapter 18. Conservation Status of the Amphibians of Brunei Darussalam

T. Ulmar Grafe and Indraneil Das

Chapter 19. Status and Conservation of Philippine Amphibians

Arvin C. Diesmos, Angel C. Alcala, Cameron D. Siler, and Rafe Brown

Chapter 20. Human Impact on Amphibian Decline in Indonesia

Djoko T. Iskandar

Chapter 21. Amphibians of Timor-Leste: a Small Fauna under Pressure.

Hinrich Kaiser, Mark O’Shea, and Christine M. Kaiser

Chapter 22. Status and Diversity of the Frogs of New Guinea

Allen Allison

PART 2. STATUS OF CONSERVATION AND DECLINE OF AMPHIBIANS: NORTHERN AFRICA (EDITED BY STEPHEN D. BUSACK AND HAROLD HEATWOLE), 2013. BASIC AND APPLIED HERPETOLOGY, ASOCIACIÓN HERPETOLÓGICA ESPAÑOLA, MADRID. PAPERBACK.

Chapter 23. Introduction.

Harold Heatwole and Stephen D. Busack

Chapter 24. Amphibian Conservation in Mauritania

José Manuel Padial, Pierre-André Crochet, Philippe Geniez, and José Carolos Brito

Chapter 25. Amphibians of Morocco, including Western Sahara: A Status Report

Ricardo Reques, Juan M. Pleguezuelos, and Stephen D. Busack

Chapter 26 Diversity and Conservation of Algerian Amphibian Assemblages

José A. Mateo, Philippe Geniez, and Jim Pether

Chapter 27. Conservation Status of Amphibians in Tunisia

Nabil Amor, Mohsen Kalboussi, and Khaled Said

Chapter 28. Amphibians in Libya: A Status Report

Adel A. Ibrahim

Chapter 29. Amphibians of Egypt: A Troubled Resource

Adel A. Ibrahim

Chapter 30. Withdrawn

PART 3. STATUS OF CONSERVATION AND DECLINE OF AMPHIBIANS: WESTERN EUROPE (EDITED BY HAROLD HEATWOLE AND JOHN W. WILKINSON), 2013. PELAGIC PUBLISHING, EXETER, UK. PAPERBACK.

Chapter 31. Infectious Diseases that May Threaten Europe’s Amphibians

Trent W. J. Garner, An Martel, Jon Bielby, Jaime Bosch, Lucy G. Anderson, Anna Meredith, Andrew A. Cunningham, Matthew C. Fisher, Daniel A. Henk, and Frank Pasmans

Chapter 32. Conservation and Declines of Amphibians in Ireland

Ferdia Marnell

Chapter 33. Amphibian Declines and Conservation in Britain

John W. Wilkinson and Richard A. Griffiths

Chapter 34. Conservation and Declines of Amphibians in The Netherlands

Anton Stumpel

Chapter 35. Amphibian Declines and Conservation in Belgium.

Dirk Bauwens and Gerald Louette

Chapter 36. Amphibian Declines and Conservation in France

Jean-Pierre Vacher and Claude Miaud

Chapter 37. Conservation and Declines of Amphibians in Spain

Cesar Ayres

Chapter 38. Conservation and Declines of Amphibians in Portugal

Rui Rebelo, Maria José Domingues Castro, Maria João Cruz, José Oliveira, José Teixeira, and Eduardo Crespo

PART 4. STATUS OF CONSERVATION AND DECLINE OF AMPHIBIANS: SOUTH-EASTERN EUROPE AND TURKEY (EDITED BY HAROLD HEATWOLE AND JOHN W. WILKINSON), 2015. PELAGIC PUBLISHING, EXETER, UK. PAPERBACK.

Chapter 39. The Amphibians of the Italian Region: a Review of Conservation Status

Franco Andreone

Chapter 40. Amphibian Conservation and Declines in Malta

Patrick J. Schembri

Chapter 41. Conservation and Declines of Amphibians in Croatia

Olga Jovanovic and Dušan Jelić

Chapter 42. Conservation and Declines of Amphibians in Slovenia

David Stanković, Martina Lužnik, and Katja Poboljšaj

Chapter 43. Conservation and Decline of European Amphibians: The Republic of Serbia

Jelka Crnobrnja-Isailović and Momir Paunović

Chapter 44. Amphibian Declines and Conservation in Montenegro

Ruža Ćirović

Chapter 45. Amphibian Declines and Conservation in Bosnia-Herzegovina

Avdullahu Adrović

Chapter 46. Conservation and Protection Status of Amphibians in Macedonia

Bogoljub Sterijovski

Chapter 47. Amphibians of Albania

Idriz Haxhiu

Chapter 48. Declines and Conservation of Amphibians in Greece

Konstantinos Sotiropoulos and Petros Lymberakis

Chapter 49. Amphibian Conservation and Decline in Romania

Dan Cogălniceanu and Laurenţiu Rozylowicz

Chapter 50. Conservation and Decline of Amphibians in Hungary

Judit Vörös, István Kiss, and Miklós Puky

Chapter 51. Conservation and Declines of Amphibians in Bulgaria

Nikolay Dimitrov Tzankov and Georgi Sashev Popgeorgiev

Chapter 52. Amphibian Conservation and Decline in Turkey

Kurtuluş Olgun and Nazan Taşkın Üzüm

Chapter 53. Conservation of Amphibians in Cyprus

Petros Lymberakis, Haris Nicolaou, and Konstantinos Sotiropoulos

PART 5. STATUS OF CONSERVATION AND DECLINE OF AMPHIBIANS: NORTHERN EUROPE (EDITED BY HAROLD HEATWOLE AND JOHN W. WILKINSON. PELAGIC PUBLISHING, EXETER, UK, IN PROGRESS.

Chapter 54. Status and Conservation of Amphibians in Luxembourg

Laura Wood, Edmee Engel, Richard A. Griffiths, Roland Proess, and Laurent Schley

Chapter 55. Germany (in progress)

Richard Podloucky and Andreas Nöllert

Chapter 56. Conservation and Declines of Amphibians in Poland

Maciej Pabijan and Maria Ogielska

Chapter 57. Amphibian Conservation in Switzerland

Benedikt R. Schmidt and Silvia Zumbach

Chapter 58. Amphibian Declines and Conservation in Austria

Marc Sztatecsny

Chapter 59. Conservation and Decline of European Amphibians: The Czech Republic

Lenka Jeřábková, Martin Šandera, and Vojtech Baláž

Chapter 60. Amphibian Declines and Conservation in Slovakia

Ján Kautman and Peter Mikuliček

Chapter 61. Conservation Status of Amphibians in Norway

Leif Yngve Gjerde

Chapter 62. Conservation Measures and Status of Amphibians in Sweden

Claes Andrén

Chapter 63. Decline and Conservation of Amphibians in Finland

Ville Vuorio and Jarmo Saarikivi

Chapter 64. Decline and Conservation of Amphibians in Estonia

Riinu Rannap

Chapter 65. Decline and Conservation of Amphibians in Latvia

Aija Pupina, Mihails Pupins, Andris Ceirans, and Agnese Pupina

Chapter 66. Lithuania (in progress)

Giedrius Trakimas

Chapter 67. Denmark

Kåre Fog, Lars Christian Adrados, Andreas Andersen, Lars Briggs, Per Klit Christensen, Niels Damm, Finn Hansen, Martin Hesselsøe, and Uffe Mikkelsen

Contributors to Part 6 (Australia, New Zealand, and Pacific Islands)

CO-EDITORS (ALSO AUTHORS)

HEATWOLE, Harold, Department of Zoology, The University of New England, Armidale, New South Wales 2351, Australia, and Department of Biology, North Carolina State University, Raleigh, NC 27695-7617, USA. [e-mail]

ROWLEY, Jodi J. L., Australian Museum Research Institute, Australian Museum, 1 William Street, Sydney, New South Wales 2010, Australia, and Centre for Ecosystem Science, Biological, Earth and Environmental Sciences, University of New South Wales, Sydney, Australia. [e-mail]

AUTHORS

ALFORD, Ross A., College of Marine and Environmental Science, Centre for Tropical Biodiversity and Climate Change, James Cook University, Townsville, Queensland 4811, Australia. [e-mail]

BELL, Ben D., Centre for Biodiversity & Restoration Ecology, School of Biological Sciences, Victoria University of Wellington, PO Box 600, Wellington 6140, New Zealand. [e-mail]

BISHOP Phillip J., Department of Zoology, University of Otago, PO Box 56, Dunedin 9054, New Zealand. [e-mail]

BRADFIELD, Kay, Perth Zoo, PO Box 489, South Perth, Western Australia 6951, Australia. [e-mail]

BYRNE, Philip, Biological Sciences, University of Wollongong, Northfields Avenue, Wollongong, New South Wales 2522, Australia. [e-mail]

CLEMANN, Nick, Arthur Rylah Institute for Environmental Research, Victorian Department of Environment, Land, Water and Planning, Heidelberg, Victoria 3084, Australia. [e-mail]

COGGER, Harold G., Australian Museum Research Institute, Australian Museum, 1 William Street, Sydney, New South Wales 2010, Australia. [e-mail]

COOTE, David, New South Wales Office of Environment and Heritage, PO Box 2111, Dubbo, New South Wales 2830, Australia. [e-mail]

EVANS, Murray, ACT Government, Level 1 Building 3, 9 Sandford St, Mitchell, Australian Capital Territory 2911, Australia. [e-mail]

FISHER, Robert N., U.S. Geological Survey, Western Ecological Research Center, San Diego Field Station, 416 Spruance Road Suite 200, San Diego, CA 92101-0812, USA. [e-mail]

GILBERT, Deon, Melbourne Zoo, Elliott Avenue, Parkville, Victoria 3052, Australia. [e-mail]

GILLESPIE, Graeme R., Flora and Fauna Division, Department of Environment and Natural Resources, PO Box 496, Palmerston, Northern Territory 0831, Australia, and School of Biosciences, University of Melbourne, Parkville, Victoria 3010, Australia. [e-mail]

HOLLIS, Greg, Baw Baw Shire Council, 90 Smith St, Warragul, Victoria 3820, Australia. [e-mail]

HUNTER, David, New South Wales Office of Environment and Heritage, PO Box 1040, Albury, New South Wales 2640, Australia. [e-mail]

LEMCKERT, Frank, SMEC (Member of the Surbana Jurong Group), Level 5, 20 Berry Street, North Sydney, New South Wales 2060, Australia, and Australian Museum Research Institute, Australian Museum, 1 William Street, Sydney, New South Wales 2010, Australia. [e-mail]

MAHONY, Michael, School of Environmental and Life Sciences, The University of Newcastle, University Drive, Callaghan, New South Wales 2308, Australia. [e-mail]

MARANTELLI, Gerry, Amphibian Research Centre, PO Box 1365, Pearcedale, Victoria 3912, Australia. [e-mail]

McFADDEN, Michael, Herpetofauna Department, Taronga Conservation Society Australia, Bradleys Head Road, Mosman, New South Wales 2088, Australia, and School of Biological Sciences, University of Wollongong, New South Wales 2522, Australia. [e-mail].

NEWELL, David, School of Environment, Science, and Engineering, Southern Cross University, PO Box 157, Lismore, New South Wales 2480, Australia. [e-mail]

OCOCK, Joanne, Water, Wetlands and Coastal Science, New South Wales Office of Environment and Heritage, Sydney, New South Wales, Australia. [e-mail]

PHILIPS, Annie, Natural and Cultural Heritage Division, Department of Primary Industries, Parks, Water and Environment, 134 Macquarie Street, Hobart, Tasmania 7000, Australia. [e-mail]

ROBERTS, J. Dale, Centre of Excellence in Natural Resource Management, University of Western Australia, PO Box 5771, Albany, Western Australia 6332, Australia. [e-mail]

SCHEELE, Ben, Fenner School of Environment and Society, Australian National University, Canberra, Australian Capital Territory 2601, Australia. [e-mail]

SHINE Richard, Heydon-Laurence Building A08, School of Life and Environmental Sciences, University of Sydney, New South Wales 2006, Australia. [e-mail]

WASSENS, Skye, Institute of Land Water and Society, School of Environmental Sciences, Charles Sturt University, Albury, New South Wales 2640, Australia. [e-mail]

WEST, Matt, School of Ecosystem and Forest Science, University of Melbourne, Parkville, Victoria 3052, Australia. [e-mail]

ZUG, George, Department of Vertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Washington, DC 20013-7012, USA. [e-mail]

Preface

The late 20th century and the early 21st century has been characterised by an unprecedented deterioration of the environment of the Earth and the throes of one of the major extinction events of all time. Unmitigated deforestation, desertification, erosion and salinisation of soil, pollution of water and air, and thinning of the UV-protective ozone layer constitute dire ecological threats for life on the planet. Fossil carbon is being returned to the atmosphere at an accelerated rate with a concomitant change in the Earth’s climate that is making serious inroads into ecological stability.

The human population now exceeds the long-term carrying capacity of the Earth and subsists at its present level only because it is sustained by fossil resources of energy, soil, fresh water (laid down in aquifers in the Pleistocene) and even oxygen (produced by photosynthetic organisms over a few billion years). With continuing decrease in biodiversity, progressive destruction of essential habitats, degradation of major ecosystems, and contamination of life-support systems, it is likely that the carrying capacity of the Earth will decline below present levels – while at the same time the human population continues to rise exponentially. The outstripping of even the fossil resources, predicted to occur within the present century, presents a bleak outlook for our own species. We may well become victims ourselves of this most recent mass extinction.

While it is undisputable that many aspects of environmental degradation and loss of biodiversity is directly attributable to unwise human activities, other aspects are deemed to result from natural forces beyond the control of mankind. It is important to be able to distinguish between the two, so that attention can be focused on mitigating those effects over which we do have control. It is important to ascertain the causes of particular declines and extinctions as soon as possible, so that steps can be taken to preserve what diversity we can.

Amphibians, by virtue of their thin, moist, permeable skins, are poorly protected from harsh environments and are especially susceptible to chemical changes, desiccation, disease, and to alteration of their habitat. The biphasic lifestyle of most amphibians also exposes them to threats both on land and in water. Accordingly, it is not surprising that they manifest proportionately high extinction rates and more severe declines than do most other organisms. They are especially important to study because they serve as an early warning system portending changes that may soon impinge upon more resistant species, including ourselves, and because they are an important part of many ecosystems: driving flows of nutrients and the transfer of energy.

The series Amphibian Biology, as its name implies, treats the entire biology of amphibians, but because of the extreme importance of the decline and conservation of this taxon, four volumes are devoted to these topics. Volumes 8 and 10 contain a topical elucidation of the myriad factors responsible for amphibians’ demise and an assessment of measures that can be taken to conserve them. Volume 9 (Western Hemisphere) and 11 (Eastern Hemisphere) assess the worldwide status of the decline, extinction, and conservation of amphibians in several Parts (see Table of Contents) organised on a country-by-country basis; the present one (Volume 11, Part 6) deals with Australia, New Zealand, and the Pacific Islands.

Although the various Parts of Volume 11 have been produced by different publishers, the chapters are numbered consecutively across the entire volume. CSIRO Publishing, however, required that the present Part be numbered separately as Chapters 1–14. Consequently, in the Table of Contents that sequence is followed (in parentheses) by the corresponding designation for Part 6 (Chapters 68–81) relating to consecutive numbering across the entire volume.

The taxonomic nomenclature of amphibians is in a state of flux and is controversial. In this book, we retain use of the generic names Litoria and Cyclorana (following the taxonomy of AmphibiaWeb, http://amphibiaweb.org) pending a comprehensive resolution of phylogenetic relationships.

There are various websites that estimate the current vulnerability of amphibian species to extinction and the present status of their decline. These websites change as status changes. Amphibian Biology does not compete with those, but rather is a ‘time capsule’ providing a perspective by present-day experts on the conservation of amphibians and the challenges faced by this taxon in each Australo-Pacific region. Because species are declining or going extinct while chapters are being written and put to press, in one sense the present treatise already will be out of date by the time it is published. In another sense, however, it is timeless because one can return to it for information on status at a specific time as a means of assessing subsequent extents and rates of change.

1Introduction

Harold Heatwole and Jodi J. L. Rowley

HISTORY OF THE ENVIRONMENTS AND BIOTA

Australia

Australia’s tectonic plate separated from the Antarctic remnant of Gondwanaland sometime before 55 million years ago and drifted northwards to abut on the Asian plate toward the end of the Oligocene about 25 million years ago (Hall 2002, 2009). During the more than 30 million years for that traverse to occur, the continent was increasingly isolated from floral and faunal exchange with other continents, except possibility for exceedingly rare overwater dispersal of waifs. Amphibians, being highly sensitive to salt water, are poor subjects for such long-distance, overwater dispersal and consequently the present batrachofauna of Australia either originated in Gondwanaland in ancient times and adaptively radiated during the long voyage northward, or arrived much more recently after the Australian and Asian plates collided (Tyler and Lee 2006). Ranidae clearly belongs to the latter category because Papurana damaeli, the sole representative in Australia of this otherwise widespread family, only occurs on Cape York Peninsula and in eastern Arnhem Land. The Australian microhylids also may have originated from an Asian source (see Chapter 2), but the remaining taxa of Australian frogs probably descended from ‘hitchhikers’ aboard ‘Ark Australia’ during its northward journey from Gondwanaland.

During this long passage, the Australian climate and vegetation varied dramatically, with an overall trend toward increasing aridity and an adaptation of some frogs to dry conditions (see Chapter 2). This trend was not a straight-line progression, however, because there were reversals superimposed on overall drying (Fujioka and Chappell 2010). It is likely that there was a pulsation of expanding aridity outward from the centre, leaving coastal pockets of moist habitat and divergence of the isolated populations contained in them, followed by a retreat of aridity, with humid regions reconnecting and the ranges of the newly formed species expanding geographically (Chapter 1 in Heatwole and Taylor 1987). The repeated cycles of speciation of isolated populations followed by expansions of range, led to radiation into a wide variety of habitats and to a few species with unusual adaptations to aridity (see Chapter 2).

The topography of Australia is one of generally low relief, with the Great Dividing Range skewed toward the eastern coast; the climatic pattern is of a monsoonal tropical north, a moist eastern and south-western temperate periphery, and an arid (desertified) core. These topographic and climatic features are reflected in the zonation of vegetation underlying the topical organisation of chapters in this book.

New Zealand

About 83 million years ago, a fragment of continental crust about the size of modern India rotated away from the Australian section of Gondwanaland to form a land mass known as Zealandia (Goldberg et al. 2008). Subsequently, the rift separating Zealandia from Australia widened to form the Tasman Sea and, over about 60 million years, Zealandia thinned and sank until only a much smaller remnant, the present New Zealand, remained above water. A boundary collision of the plate in the late Oligocene gave rise to topographic upheaval to form the Southern Alps.

New Zealand’s biota consists of a mixture of ancient Gondwanan elements and more recent arrivals (Goldberg et al. 2008). In line with their physiological characteristics, frogs do not disperse over salt water easily and, not surprisingly, New Zealand’s frogs are of Gondwanan origin. In fact, they are among the most primitive of frogs, having diverged from more modern taxa in about the mid-Triassic (see Chapter 13). All are endemic to New Zealand and, in keeping with the mesic climate there, are denizens of forests.

The oceanic islands of the Pacific

Oceanic islands are those that arise de novo from the sea without any present or previous direct connection to a continent or isolated continental fragment (such as New Zealand). Chapter 14 and Neall and Trewick (2008) summarise the tectonic history of the Pacific Basin and the islands lying within it, and that will not be detailed here. The salient point is that most oceanic islands of the Pacific are devoid of amphibians, probably because of their great distances from a continental source of terrestrial fauna and the difficulty with which amphibians disperse across salty water. Only three island-groups near the periphery of the Pacific, and hence nearer sources of continental faunas, have native frogs – the Solomon Islands, Palau, and Fiji – although some others have species introduced by humans (see Chapter 14).

THE GLOBAL CONTEXT OF THE DEMISE OF AMPHIBIANS

Throughout geologic history there have been periodic mass extinctions in which the biota of the Earth was reduced cataclysmically (McElwain and Punyasena 2007). The causes of these events have been varied: one perhaps being due to changing patterns of oceanic currents and climatic disruptions resulting from the separation of the supercontinent Pangaea into two, and eventually all seven, of the present continents. The most famous mass extinction, however, was the one marking the transition from the Mesozoic Era to the Cenozoic about 66 million years ago, attributed to the collision of a comet with the Earth, and highlighted by the precipitous disappearance of dinosaurs as well as many other taxa (Alvarez et al. 1980). The eventual fate of all species, of course, is to disappear, either through dying out completely (extinction), or by loss of the ancestral form via its transition into new, and different, taxa (evolution, speciation). Previous mass extinctions were followed by a regenerative cycle as the surviving taxa adaptively radiated into ecological niches left vacant by the demise of victims of the extinction event. Recovery requires millions of years (McElwain and Punyasena 2007).

Currently, almost one-third of all amphibian species are threatened with extinction, making them the most threatened group of terrestrial vertebrates (IUCN 2017). Thirty-three amphibian species are officially listed as recently extinct, but this likely is a severe underestimate. The causes of amphibian declines and extinctions are multiple (Heatwole 2013) and form a complex network that is nearly intractable to solution by virtue of the labyrinth of a large number of interactive links (Plate 1.1). Amphibian decline is global, but is generated by different combinations of factors in different regions, although some causes are common and widespread geographically.

One of the biggest obstacles to halting the decline of amphibian species is a lack of knowledge. At the most basic level, we still do not know how many species of amphibians there are; in the past decade, an average of 157 new species of amphibians have been described annually (AmphibiaWeb 2017). This trend shows no sign of slowing. For the species of which we are aware, we often lack information necessary to carry out informed conservation. Indeed, 24% of all currently assessed amphibian species (84% total known amphibian species to date; AmphibiaWeb 2017) are so poorly understood that their conservation status cannot be determined (IUCN 2017).

The mass extinction now in progress differs from previous ones by virtue of its largely anthropogenic origins – we are the root of most of the causes. This aspect, however, does have a positive side: there is an intelligent, sentient species involved in the event. If we can cause declines that lead to extinction, perhaps we also can devise ways to prevent, or at least ameliorate, them. Consequently, this book has two goals: (1) to contribute to an understanding of this complex phenomenon; and (2) to stimulate research into finding solutions to prevent the occurrence of the worst-case scenario. A lot of research toward these ends already has taken place and has been reviewed (see Preface); the material presented here is by way of application specifically to Australia, New Zealand, and the Pacific Islands.

With such a complex set of causes as illustrated in Plate 1.1, the tracing and quantifying of the links in the network of interactions is a formidable task. We must be vigilant to avoid facile oversimplifications. Too often, once a single correlation has been established, it is accepted as denoting a cause and effect relationship, with that conclusion becoming widely accepted and without delving into the possible intervention of other factors. The oversimplified version may then be enshrined as truth for all occasions in the lexicon of scientific mythology, and the complete suite of reasons bypassed by future investigators. Enigmas are important because they help identify such situations and allow greater focus on reconciling apparent contradictions. For example, Lane and Burgin (2008) reported that at lower elevations in the Greater Sydney region the diversity of amphibians was lower in more urbanised and polluted sites than in natural, less polluted ones, as one would expect, but that at higher elevations, the reverse occurred. In view of the complexity of possible interactions illustrated in Plate 1.1, paradoxes like this are important and should be followed up for verification and further scrutiny, rather than merely dismissed as aberrant. Resolution of such mysteries may lead to a far better understanding than could be achieved merely by unduly emphasising the more predicable outcome.

A common misconception that also tends to disguise the dynamics of natural systems is an optimistic view of their regenerative capacity. It is true that biological systems often exhibit fluctuations around some mean level, and that departures from that level bring into play forces tending to return the system toward the mean, whether the departure is above the mean or below it. The mechanisms directing such returns toward a stable equilibrium collectively are known as negative feedback, and the entire process of oscillation around a set value is called homeostasis. Regulation of body temperature in mammals is an example of such a system. When the body begins to get too hot, certain automatic responses, such as panting, sweating, or seeking shade, occur that tend to cool the body. When the body begins to cool below the equilibrium level, other responses, such as shivering, come into play and raise the body back toward the ‘normal’ level. Such mechanisms are frequent in biological systems and operate at various levels, ranging from the physiology of an individual (as in the above example) to the dynamics of biotic communities and ecosystems. This generality has inspired unwarranted optimism that homeostasis will operate consistently and that, when disturbed, ‘nature’ will automatically restore itself to its original condition, merely if left alone. That is not always the case and the more severe the disturbance, the more likely it is that some non-equilibrium state will prevail or that a new, less desirable, equilibrium will be reached (Rohde et al. 2013). In the worst-case scenario, an irretrievable situation, such as extinction, may alter the equilibrium. For example, amphibian population declines in Central America in the 1990s has resulted in large-scale ecosystem-level effects in stream habitats that persist today (Whiles et al. 2006, 2013). There are limits to homeostasis.

AMPHIBIANS IN AUSTRALIA, NEW ZEALAND, AND THE PACIFIC ISLANDS

Australia has a relatively well-known frog fauna but much remains unknown, even with respect to the true number of species. There are currently 240 native species of frogs known from Australia (see Chapter 2 for a summary of the diversity, distributions, and conservation status), but in the past decade alone, 21 new species, representing 9% of Australia’s known frog fauna, have been discovered. Most of these species were hidden within known ‘species’ that were found to be complexes of multiple, morphologically similar species (e.g. Mahony et al. 2006; Anstis et al. 2016; McDonald et al. 2016), but others have been discovered as the result of surveys in remote or previously unsurveyed areas (e.g. Hoskin and Aland 2011). The discovery of previously undescribed species of frogs in Australia is ongoing, particularly in tropical northern regions. The frog fauna of the Pacific Islands is considerably less well known (see Chapter 14), with just over 30 known species and new species found on almost every expedition (e.g. the Solomons). New Zealand’s native frog fauna currently consists of four species belonging to a single relatively ancient lineage (see Chapter 13).

Although geographically proximate, each region presents unique challenges and opportunities in amphibian research and conservation. Whereas population declines of frogs are well documented, and research and conservation have a long history in Australia and New Zealand (see Chapters 3–9, 11), present knowledge is insufficient even to determine to what extent population declines have occurred in most species (see Chapter 14).

It is hoped that this book, by presenting and critically interpreting the body of available information on the current status of amphibians in Australia, New Zealand, and the Pacific, will form the basis for future research into an urgent ecological problem. Equally important, or perhaps more so, it also is hoped that persons involved in formulating policy or executing conservation actions will be able to do so with greater understanding and success as a result of access to the accumulated knowledge and understanding summarised in this modest volume.

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