Table Of ContentTERRESTRIAL ISOPOD BIOLOGY
CRUSTACEAN ISSUES
9
General editor:
FREDERICK R. SCHRAM
Instituut voor Systematiek en Populatiebiologie, Amsterdam
A. A. BALKEMA / ROTTERDAM / BROOKHELD /1995
TERRESTRIAL
ISOPOD BIOLOGY
Edited by
M.A. ALIKHAN
Laurentian University, Sudbury, Ontario
A. A. BALKEMA / ROTTERDAM / BROOKFIELD/1995
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Table of contents
Preface VH
Introduction IX
Comparative study of metal bioaccumulation, and oxygen and nitrogen metabolism
in three terrestrial isopod species from contaminated and uncontaminated sites in
Northeastern Ontaria
MAAlikhan 1
Comparative anatomy and functional aspects of the digestive system in amphibious
and terrestrial isopods (Isopoda: Oniscidea)
J.Strus, D. Drobne & P. Licar 15
Water vapour absorption and ammonia volatilization: Adaptations for terrestriality
in isopods
Jonathan C Wright & Michael J. O 'Donnell 25
Ligia: A prototypal terrestrial isopod
Thomas H Carefoot & Barbara K Taylor 47
Mechanism and function of turn alternation mArmadillidium vulgare (Latreille)
Tang Bo-Ping, Gui Hong & Wu Tian 61
Reproduction in woodlice: Flexibility to maximise individual fitness
J. M. Dangerfield &S.R. Telford 69
Isopod distribution at different scaling levels
£ Homung &M.R Warburg 83
Diversity and spatial community structure of terrestrial isopods (Isopoda, Oniscidea)
in a mosaic of plant assemblages
Katalin Sddvecz 97
VI Preface
Effects of temperature and photoperiod on the breeding patterns of two isopod
species
M R. Warburg & Dorit Weinstein 107
The effect of leaf litter, microorganisms and Collembola on the food allocation of
Oniscus asellus (Isopoda)
MUesbeck&WTopp 121
Microhabitat selection by Armadillidium vulgare in a riparian forest: Lack of
apparent influence by leaf litter food quality
F.Heinzelmann, C S. Crawford, M. C Molles, Jr.&MR. Warburg 133
Population genetics of Armadillidium vulgare in Europe and North America
R. L Garthwaite, R. Lawson & C Sassaman 145
Contributors 201
Index 203
Preface
In 1967, Dr. Dorothy E. Bliss of the American Museum of Natural History at New
York, brought together for the first time at the New York meeting of the American
Association for the Advancement of Science scientists with an interest in terrestrial
isopods. Since then, three international symposia dealing with the 'Biology of
Isopods,' the most successful of the land colonizing crustaceans, have been held in
London (United Kingdom), Urbino (Italy), and Poitiers (France). The London meet-
ing was sponsored by the Zoological Society of London, and its proceedings contain-
ing 21 papers, were edited by Stephen Sutton and David Holdich (Zoological Society
of London Symposia 53, 1984). Proceedings of the second symposium, sponsored by
Consiglio Nazionale delle Ricerche Unione Zoologica Italiana, containing 26 presen-
tations, were published by Franco Ferrara, Roberto Argano, Claudio Manicastri,
Helmut Schmalfuss, and Stefano Taiti [Monitore zoologico italiano (N.S.) Mono-
grafia 4, 1989]. The third meeting was financed by 'Faculte des Sciences de 1'Uni-
versite de Poitiers,' 'Conseil scientifique de 1'Universite de Poitiers,' 'Conseil Gene-
ral de la Vienne' and 'Muncipalite de la Ville de Poitiers, France.' The volume,
containing 26 papers and 12 posters presented during the meeting, was organized by
Pierre Juchault and Jean Pierre Mocquard.
The present symposium was proposed for the 1992 meeting of the American So-
ciety of Zoologists by Professor Michael R. Warburg of Technion, Israel Institute of
Technology at Haifa, during his stay during 1991 with me at the Laurentian Univer-
sity. This explains my involvement in the organization of the meeting, and the edit-
ing of its proceedings. The sponsorship was provided by The Crustacean Society.
Unfortunately, because of the timing of the meeting, fewer than 20 colleagues from
various parts of the world were able to attend. Nevertheless, more than twelve papers
and four posters on various aspects of the biology of isopods were presented both
during the symposium and the general session. Out of these, ten papers are being
published in this volume in addition to some other contributions on the subject that
were not part of the original symposium. The paper presented by Barbara Taylor and
T. H. Carefoot on the 'Evolution of terrestrial life in isopods: loss of gas exchange &
survival ability in water,' has been accepted for publication in the Canadian Journal
of Zoology (1993).
I am grateful to Drs. Robert W. Elner and Richard Brusca of The Crustacean So-
ciety for their continued encouragement, to Dr. Thomas H. Carefoot of the Depart-
vn
VIII Preface
ment of Zoology of the University of British Columbia, Dr. Cliff Crawford of the
Biology Department of the University of New Mexico, and Dr. Michael Warburg of
the Department of Biology at the Technion at Haifa, for their support and help in the
organization of the symposium, to Dr. Stephano Taiti of Centra di Studio per la
Faunistica ed Ecologia Tropicali, CNR, Florence (Italy) for providing me with the
addresses of various scientists and sending out the notice of the meeting, to the
American Society of Zoologists for the waiver of registration fee for the participants
in the symposium, and to my various colleagues, who, because of the holiday season
made personal sacrifice and attended the meeting. Last of all my thanks to Bridget
Farley of the American Society of Zoologists for her support, advice and assistance.
She deserves a part of the credit for the success of this symposium.
M.A. Alikhan
Sudbury, Canada
June 11, 1993
Introduction
Last time isopods were discussed on American soil was 25 years ago at the sympo-
sium on 'The terrestrial adaptations of Crustacea,' arranged by Dorothy Bliss and
Eric Edney during the AAAS meeting in New York, December 1967. Meanwhile the
isopodologists held three symposia in Europe where most of the research on this
group was conducted. All of the symposia volumes have been published. One aim of
this meeting was to stimulate research on American isopods some of which are en-
demic species and very little is known about them.
This fascinating group is the only crustacean group with representatives in all ter-
restrial ecosystems ranging from the sea shore to the desert. Several species have
colonized deserts, some of them are highly successful and abundant. Thus, this group
can be considered as a model for the successful transition on land.
Much of the early work was descriptive concerning structure. Other early work
dealt with ecological and zoogeographical aspects. More recent research dealt with
physiological adaptations related to water and thermal balance. In recent years, more
attention was devoted to ecological studies dealing with reproductive strategies, and
population and community structure.
The symposium deals with two main subjects: 1) The effects of stressful condi-
tions on the individual animal as reflected by its survival, or by the disruption of its
normal reproductive patterns. 2) The distribution of the isopods and their selection of
microhabitat. Thus, one paper by Strus, Drobne, and Licar dealt with the comparative
anatomy of the digestive system in two amphibious and two terrestrial isopod spe-
cies, while another paper by Alikhan discusses the effect of metal pollutants on the
metabolism. A third paper by Wright and O'Donnell describes the ways ammonia is
excreted by a terrestrial isopod inhabiting (Porcellio) a mesic habitat. Very little is
known about excretion in oniscid isopods, in particular of interest are the desert spe-
cies because of their water shortage problem. Another paper by Heinzelmann, Craw-
ford, Warburg, and Molles dealt with the effects of temperature and photoperiodism
on the reproductive pattern in two isopod species: Armadillo officinalis and Schizid-
ium tiberianum, while Dangerfield and Telford discussed the importance of life his-
tory variation with regards to reproductive tactic. The second subject was discussed
in three lectures. One paper by Szlavecz dealt with the distribution of isopod species
in an old park in Europe. Warburg and Weinstein dealt with the distribution of
isopods as related to stone coverage in a Mediterranean grassland. Finally, a third
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