Table Of ContentAmphibian Biology
Edited by
Harold Heatwole
Volume 8
Amphibian Decline: Diseases,
Parasites, Maladies and Pollution
Co-editor for this volume
John W. Wilkinson
Published by
Surrey Beatty & Sons
This book is copyright. Apart from any fair dealing for purposes of
private study, research, criticism or review, as permitted under the
Copyright Act, no part may be reproduced by any process without
written permission. Enquiries should be made to the publisher.
Surrey Beatty & Sons Pty Ltd is a member of Copyright Agency
Limited (CAL) in Australia and is a respondent to overseas collecting
societies.
The National Library of Australia
Cataloguing-in-Publication Entry:
Amphibian Biology. Volume 8, Amphibian Decline:
Diseases, Parasites, Maladies and Pollution
Includes Index.
ISBN 0 949324 53 1 (set)
ISBN 0 949324 54 X (v. 1)
ISBN 0 949324 60 4 (v. 2)
ISBN 0 949324 72 8 (v. 3)
ISBN 0 949324 87 6 (v. 4)
ISBN 0 949324 94 9 (v. 5)
ISBN 0 949324 95 7 (v. 6)
ISBN 978 0 98031 13 10 (v. 7)
ISBN 978 0 98031 13 34 (v. 8)
1. Amphibians - Classification. I. Heatwole, Harold.
Published 2009
PRODUCEIND A USTRALBIYA
SURREY BEATTY & SONS PTY LIMITED
PO Box 8159, Baulkham Hills NSW 2 153 Australia
Preface to Series
T
ERE are several outstanding treatises of amphibian biology. Biology of Amphibians
(Duellman and Trueb 1986) is an excellent general work, clearly written and well
illustrated, and with a remarkable depth and breadth for a single volume. It will be the
standard general reference on amphibians for many years to come and the present
generation of herpetologists will consider it the "amphibian bible" much as their
predecessors regarded G. K. Noble's (1931) The Biology of the Amphibia. No single volume,
however, fulfils the need for a sequential, monographic treatment of specialized topics.
A few individual subjects have been treated in considerable depth. These include the three
volumes of Physiology of the Amphibia (Moore 1964; Lofts 1974, 1976), Frog Neurobiology, a
Handbook (Llinas and Precht 1976) and The Rep-oductive Bwlogy of Amphibzam (Taylor and
Guttman 1977), but it has now been many years since these appeared and many topics are in
need of updating. Environmental Physiology of the Amnphihm (Feder and Burggren 1992) did that
for some aspects and treated other new ones, but it too is now getting out of date. Two books,
Pattems of Dish'bution of Amphibians (Dueban 1999) and Tadpoles (McDiarmid and Altig 1999)
closed out the 20h century with excellent reviews of amphibian biogeography and of the biology
of larval amphibians, respectively. Starting the millennium was a multi-volume treatment of
reproductive biology and phylogeny: Rwuctive Biology and Phylogeny of Urodela (edited by David
M. Sever) and Reproductive Biology and Phylogeny of Anura (edited by Barrie Jamieson), both in
2003, followed by Reproductive Biology and Phylogeny of Gymnophiona (edited by Jean-Marie
Exbrayat) in 2004. Reproduction was further treated by Ogielska (2009) in Reproduction of
Amphibians. Recent molecular techniques led to a rearrangement of amphibian taxonomy (Frost
et al. 2006). Behaviour and ecology of amphibians was given a recent, comprehensive, thoughtfd
update (Wells 2007). Collectively, all the above works, excellent though they are, still leave large,
unaddressed gaps in amphibian biology.
Recognizing that the discipline of amphibian biology had reached sufficient maturity
to warrant detailed, multi-volume review and that such a need had been filled only partly,
and with no commitment to continuation, the present series, Amphibian Biology, was launched
in 1994. The present volume represents the eighth in the series. Amphibian Biology does
not compete with the titles mentioned above. Topics recently reviewed elsewhere are not
covered in current volumes, but are reserved for such time as hrther update is required.
The need for this series was evidenced by the enthusiastic response from potential
authors. Of the 64 people contacted with invitations to contribute to the first few volumes,
only three declined, and then because of heavy commitments otherwise. Most expressed
the view that such a series was long overdue. With this initial encouragement the series
was launched and it is continuing to enjoy undiminished support.
Amphibian Biology was inspired by Biology of the Reptilia (1969-2008), edited by Carl Gans,
and is intended as a companion to that series. Biology of the Reptilia is a unique, monumental
contribution to herpetology and has become the most authoritative single source of
information on reptiles that is available. Comprehensive treatments of all aspects of reptilian
biology are presented in detail and are exhaustively documented by literature. It has been,
and continues to be, invaluable. It is hoped that Amphibian Biolopy will serve herpetologists
in the same way and that it will maintain the high standard set by its reptilian counterpart.
Raleigh, North Carolina, USA Harold Heatwole
January 1993, revised August 2009 Series Editor
REFERENCES
Duelhnan, W. E., 1999. "Patterns of Distribution of Amphibians". The John Hopkins University Press, Baltimore.
Duellman, W. E. and Tmeb. L., 1986. "Biology of Amphibians". McGraw-Hill Book Company, New York.
Eibrayat, J.-M. (ed.), 2004) "Reproductive Biology and Phylogeny of Gymnophiona". Science Publishers, Enfield.
Mez M. E. and Burggren, W., (ed.s) 1992. "Environmental Physiology of the Amphibians". University of Chicago
Press, Chicago.
VI AMPHIBIAN BIOLOGY
Frost, D. R., Grant, T, Faivovich, J., Bain, R., Haas, A., Haddad, C. F. B., de S., R., Channing, A., Wilkinson, M.,
Donnellan, S. C., Raxworthy, C., Campbell, J. A. , Blotto, B. L., Moler, I?, Drewes, R. C., Nussbaum, R. A., Lynch,
J. D., Green, D. M. and Wheeler, W., 2006. The amphibian tree of life. Bulletin of the American Museum of Natural
History 297: 1-370
Gans, C. (ed.), 1969-1998. "Biology of the Reptilia". 19 vols, Academic Press, New York; Alan R. Liss, Inc., New
York; The University of Chicago Press, Chicago; Society for the Study of Amphibians and Reptiles.
Jamieskon, B. G. M. (ed.), (ed.), 2003 "Reproductive Biology and Phylogeny of Anura". Science Publishers, Enfield.
Llins, R. and Precht, W., 1976. "Frog Neurobiology, a Handbook. Springer-lkrlag, Berlin.
Lofts, B. (ed.), 1974 "Physiology of the Amphibia", volume 2. Academic Press, New York.
Lofts, B. (ed.), 1976. "Physiology of the Amphibia", volume 3. Academic Press, New York.
McDiarmid, R. W. and Altig, R., 1999. "Tadpoles". The University of Chicago Press, Chicago
Moore, J. A. (ed.), 1964 "Physiology of the Amphibia". Academic Press, New York.
Noble, G. K., 1931. "The Biology of the Amphibia". McGraw-Hill Book Company, New York.
Ogielska, M. (ed.), 2009 "Reproduction of Amphibians". Science Publishers, Enfield.
Sever, D. M. (ed.), 2003 "Reproductive Biology and Phylogeny of Urodela". Science Publishers, Enfield.
Taylor, D. H. and Gutunan, S. I. (eds.), 1977. "The Reproductive Biology of Amphibians". Plenum Press, New York.
Wells, K. D. 2007. "The Ecology and Behavior of Amphibians". The University of Chicago Press, Chicago.
Preface to Volume 8
T
E late 20th century and the early 21" century has been characterized by an
unprecedented deterioration of the environment of the earth and the throes of one of
the major extinction events of all time. New diseases have emerged. Unmitigated deforestation,
desertization, erosion and salinization 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 likely to make serious inroads into ecological stability.
The human population now exceeds the long-term carrying capacity of the earth and is
able to subsist at its present level only because it is sustained by fossil resources of energy,
soil, water and even oxygen. 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. The outstripping of even its
fossil resources, likely to occur within the present century, presents a bleak outlook for our
own species. We may well become a victim ourselves of this most recent mass extinction.
While it is indisputable 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 cycles beyond the influence 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 humans
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 preseIve as much diversity as possible.
Amphibians, by virtue of their thin, moist, permeable skins, are poorly protected from
harsh environments and are especially susceptible to chemical changes, desiccation, and
alteration of habitat. 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 as they serve as an early-warning system portending changes
that may soon impinge upon more resistant species, including Homo sapzens.
The topic of conservation and decline of amphibians will be the subject of several
volumes. The present volume (8) is devoted to diseases, maladies, and parasites of
amphibians and how these relate to decline, followed by the impact of pollution, of various
kinds and from various sources, on amphibian populations. Finally, a chapter is devoted to
the phenomenon of climatic change.
Subsequent volumes will treat (1) the roles of anthropogenic influences such as habitat
change; introduction of alien species; roadkills; direct harvesting, trade, and use of amphibian
species by humans, (2) various ecological, phylogenetic, and geographic correlates of amphibian
decline, (3) monitoring programmes and conservation practices such as establishment of rehgia;
captive breeding and re-introduction; and mitigation; as well as the application of education.
The last volumes will assess the global status of conservation and decline on a region-by-region
basis to serve as a benchmark for subsequent changes that take place.
There is modest overlap in subject matter among chapters. This is deliberate as particular
aspects interact and it is more convenient to have complete coverage of a given topic without
having to shift back and forth between chapters. Each chapter was designed to stand alone.
Many of the chapters of these volumes were in mid-preparation when the monograph
by Frost et al. (2006) appeared. The new taxonomic arrangements and nomenclatural
changes espoused by Frost et al. are still controversial and there has been no editorial policy
demanding authors either to adhere to it, or not to do so. Consequently, some chapters
have the older scheme and others the newer one, as the authors' own assessments dictated.
Raleigh, North Carolina, USA Harold Heatwole
17 January 2009 Series Editor
VIII AMPHIBIAN BIOLOGY
Lee Bergel
Dedication
I? RTUNATELY for amphibians Lee Berger decided to take a year off from her veterinary
studies in 1991 to travel around Australia to see what research on wildlife diseases was
being conducted. On that trip she met the eclectic scientist, veterinarian and doctor Rick
Speare in Townsville, Queensland, and worked for him on an anti-parasitic drug that was
causing mortality in wallabies. Little did she know how that would change our lives. Several
years later, in 1995, while she was working as a veterinarian in Melbourne, Victoria, I
remember her excitedly putting down the telephone to tell me that Rick had offered her
a PhD position to determine the spreading agent that he thought was causing frog decline
and extinction in Queensland. She moved to the Australian Animal Health Laboratory in
Geelong, Victoria, to study samples collected by Keith McDonald during a mass mortality
of frogs at Big Tableland, Queensland. This is the premier virology laboratory in Australia
and she was given the task of finding a virus in those samples. During six months of
negative findings for viruses she discovered a novel organism that we now know as
Batrachochytm'um dendrobatidis infecting the skin of these frogs as well as others submitted to
the laboratory as dead or sick. The key breakthrough came in 1996 when this pathogen
appeared in winter dieoffs and Lee decided to take this organism seriously and started to
investigate the disease it caused, chytridiomycosis, and its role in amphibian declines and
extinction.
Since the early 1980s scientists were unable to explain declines and extinctions of
amphibians in protected habitats until Lee's discovery. Her work, and that of her colleagues,
was published in the Proceedings of the National Academy of Sciences (Berger et al. 1998)
and they were awarded the Australian Commonwealth Scientific and Industrial Research
Organisation medal for science. It wasn't all smooth sailing, however. There were numerous
difficulties working with a novel pathogen and in the field of amphibian disease where there
was little expertise. She spent many late nights at work overcoming these obstacles. There
was also the pressure of solving the cause of amphibian declines as well as intense scrutiny
of her work and opposition to the hypothesis that a spreading disease was the cause of
amphibian decline. Since then, she has continued to make major contributions in
understanding the epidemiology and pathogenesis of chytridiomycosis and has investigated
other emerging diseases of amphibians. She has also devoted much of her time in assisting
others in Australia and overseas in conducting research on amphibian diseases.
Significant management recommendations for managing the disease have been
adopted based on her work. These include (1) the Amphibian Ark Progx-am that operates
an emergency captive-breeding programme to prevent extinctions of populations and
species due to outbreaks of chytridiomycosis, (2) the World Animal Health Organisation's
(OIE) listing of chytridiomycosis as a notifiable disease in order to prevent its spread,
and (3) the Australian Government listing it as a key threatening process in order to
prevent its spread and to abate its current impact on threatened species of frogs. The
spectacular informative nature of Dr. Berger's work demonstrates the risk of introducing
virulent pathogens into na'ive host populations and ultimately will lead to better
conservation of amphibians in particular and in the long term to better conservation
of wildlife in general.
Currently, Dr. Berger holds a Postdoctoral Research Fellowship at James Cook University,
balancing work on amphibian diseases with the care of three young children. It has been
my pleasure to know and work with Lee, who is not only an outstanding scientist but a
lovely human being. I am extremely pleased that this volume on amphibian conservation
is dedicated to her in recognition of how much she has contributed already to this field
during her career.
co tor
Creek, Queensland, Australia Lee F. Skerratt
