Mites: Ecology, Evolution & Behaviour: Life at a Microscale

505 Pages·2013·11.95 MB·English

by  David Evans Walter

David Evans Walter
Heather C. Proctor
Mites: Ecology,
Evolution &
Behaviour
Life at a Microscale
Second Edition
Mites: Ecology, Evolution & Behaviour

David Evans Walter (cid:129) Heather C. Proctor
Mites: Ecology, Evolution
& Behaviour
Life at a Microscale
Second Edition
David Evans Walter Heather C. Proctor
Invertebrate Zoology Biological Sciences
University of the Sunshine Coast University of Alberta
Royal Alberta Museum Edmonton, AB, Canada
Edmonton, AB, Canada
ISBN 978-94-007-7163-5 ISBN 978-94-007-7164-2 (eBook)
DOI 10.1007/978-94-007-7164-2
Springer Dordrecht Heidelberg New York London
Library of Congress Control Number: 2013948357
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Preface to the Second Edition
Most mites are minuscule, barely perceptible to even the sharpest eyes, but size is a
poor guide to importance. When a crop disappears under a blanket of silk or stored
food turns into a seething mass of hairy motes, then the numbers of mites will be
legion. If the mite is an itch mite burrowing in one’s skin or a tick or chigger inject-
ing virulent rickettsiae into one’s blood, then a single mite may be enough to affect
one’s life dramatically.
Some akari were known to the more perceptive of ancient Greeks (Kevan 1985)
and some of their effects were understood long ago (e.g. the cause of scabies has
been known since 1689), but few mites caused comment until the late 1700s (Prasad
1982). At that time, fewer than 100 species of ‘ticks’, stored-product pests and
brightly coloured plant and water mites had been discussed in the literature before
Linnaeus (Baker and Wharton 1952), and Linnaeus himself included only 29 spe-
cies of Acarus in the tenth edition of his Systema Naturae . Not until the late 1800s
was the term ‘acarologist’ coined to describe those who study mites, and it was
another 50 years before the word appeared in dictionaries (Krantz 1996).
Advances in microscopy contributed to the birth of this new science, but even
more important was the industry and imagination of acarologists such as G. Canestrini,
A.D. Michael, A. Nalepa, E. Trouessart and, in particular, Antonio Berlese in the
late 1800s. It is largely through Berlese’s work that scientists fi rst began to appreciate
the ubiquity and diversity of the Acari – and how intricately they are linked to every
aspect of terrestrial life.
Acarology prospered during its fi rst century, nurtured by the excitement of dis-
covery and by medical, veterinary and agricultural needs. In the last few decades,
however, like other important disciplines that require an understanding of whole
organisms (such as entomology and nematology), acarology has been eclipsed by a
fascination with technology, subcellular processes and genomes. As a result, no
matter how well versed they may be in cellular biology and genetics, many of
today’s students are not trained to deal with living organisms or the problems those
organisms can cause.
The repercussions of this ignorance are already being felt, especially in the pro-
liferation of pests, the spread of disease vectors and the alarming incidence of
v
vi Preface to the Second Edition
imaginary mite infestations (Delusional Parasitosis) in the general public. Still, not
all is gloomy. The transformation of molecular biology from a separate fi eld of
study to a tool that is becoming increasingly integrated into ecology, systematics
and other fi elds of evolutionary biology is a promising change. For example, there
is increasingly strong evidence from molecular studies that the Acari are diphyletic
and that ‘miteness’ has evolved independently in two relatively distantly related
lineages of arachnids. As well, molecular techniques are increasingly useful in
determining species boundaries, population structuring, feeding biology and impor-
tant life history characteristics from reproductive biology to virulence. New com-
puter technology is supporting a renaissance in interactive identifi cation tools. The
Internet has transformed our ability to retrieve and exchange information with col-
leagues around the world and to make taxonomic tools available to an international
audience. As retirement and death continue to thin the ranks of trained acarologists,
we take hope in these innovations and in the burgeoning number of young acarolo-
gists outside of North America and Europe and the continuing role of the OSU
Acarology Summer Program in training biologists in mite taxonomy and contribut-
ing to the global fellowship of acarologists.
Our goal i n revising this book is to attract new students to acarology by showing
them that mites do interesting and exciting things. To that end, we have doubled the
number of illustrations and poured over more than 1,600 scientifi c publications look-
ing for examples of why mites are interesting to study. We highlight the roles that
mites have played in the development of important theoretical concepts in ecology
and evolution (e.g. local mate competition, prey refugia, multi-level selection and
tritrophic level interactions) and also emphasize that, as Dana Wrensch has said,
‘mites do things they shouldn’t’. In many cases, the lives of mites clearly demon-
strate that currently accepted theory is fl awed. We also reveal the depth and breadth
to which mites have permeated the world – from sequoias to seamounts to sea lions –
but most of all, we strive to transmit the wonder and affection that we feel for these
most miniature of arthropods, with the hope that our acarophilia will infect others.
Quite a few biologists with at least mild cases of this affl iction have contributed
valuable suggestions after reading drafts of chapters, when reviewing earlier works
or during discussions (semi-sober and otherwise). We think it safe to say that none
of these friends and colleagues would agree with everything we say in this book, but
all have contributed to its more lucid and informative sections. We would like to
thank everyone who has provided help and support, and though it is easy to forget
one’s debts, these names spring immediately to mind: Jenny Beard, Fred Beaulieu,
Tyler Cobb, Valerie Behan-Pelletier, Andre Bochkov, Kaylee Byers, John Clark,
Matt Colloff, Rob Colwell, John Cooke, Dac Crossley, Ashley Dowling,
Doug Craig, Jacek Dabert, Jason Dunlop, Qinghai Fan, Farid Faraji, Norm Fashing,
Ray Fisher, Gordon Gordh, Bruce Halliday, Mark Harvey, Marjorie Hoy, Mark
Judson, Jim Kierans, Hans Klompen, Wayne Knee, Jerry Krantz, Evert Lindquist,
Zoë Lindo, Rogelio Macías-Ordóñez, Serge Mironov, Masha Minor, Jim McMurtry,
Helen Nahrung, Jeffrey Newton, Roy Norton, Barry OConnor, Norm Platnick, the
late Frank Radovsky, Susanne Randolf, Heinz Schatz, Owen Seeman, Matthew
Shaw, Bruce Smith, Ian Smith, Diana Wall, Gerd Weigmann, Jay Yoder and
Preface to the Second Edition vii
Zhi- Qiang Zhang. We would also like to thank Michael Caldwell for allowing us to
use his striking drawing of a rampant eurypterid and our students Catherine (née
Bryant) Harvey and Juanita Choo for their help with many of the line drawings.
Anthony O’Toole, Caroline Meacham and George Braybrook enhanced our skills at
the scanning electron microscope. We would also like to thank Zuzana Bernhart at
Springer for her editorial support. Finally, the senior author would like to express
his appreciation to the entire Nahrung family of Miva, Queensland, for their kind-
ness in allowing him to spend weeks immersed in solitude and an extraordinary
natural diversity while a large part of this second edition was taking form.
Additionally, our knowledge and appreciation of mites has grown in direct pro-
portion to our ability to study these fascinating animals. We have funded some of
our studies out-of-pocket, but that largesse has never been suffi cient to tackle more
than interesting sidelines. Fortunately, mites are major components of all terrestrial
ecosystems and there are few research projects that cannot be used to learn some-
thing exciting about mites. The senior author is especially appreciative of the sup-
port he received from the National Science Foundation (Graduate Research
Scholarship and Postdoctoral Fellowship), the USDA Agricultural Research Service
(Postdoctoral Fellowship), the Australian Research Council (Postdoctoral
Fellowship, Large and Small Grant Schemes), the Australian Biological Resources
Survey, the Department of Entomology at the University of Queensland, the Alberta
Biodiversity Monitoring Institute and the Royal Alberta Museum. The junior author
wishes to thank the Natural Sciences and Engineering Research Council of Canada,
the Department of Biology at Queen’s University, the Australian Research Council’s
Small Grant Scheme, the Australian School of Environmental Studies at Griffi th
University and the Department of Biological Sciences at the University of Alberta
for their support.
David Evans Walter
Heather C. Proctor

Contents
1 What Good Are Mites? ........................................................................... 1
What Is a Mite? ......................................................................................... 1
Why Study Mites? ..................................................................................... 5
What Follows? .......................................................................................... 7
References ................................................................................................. 9
2 The Origin of Mites: Fossil History and Relationships ....................... 11
The Cambrian Explosion and the Rise of the Arthropoda ........................ 11
The First Major Dichotomy: Mandibulata Versus Chelicerata ................. 15
A Review of Arthropod Limb Structure, Metamerism and Tagmosis ...... 18
Marine Euchelicerates ............................................................................... 20
Scorpionida: The First Arachnids? ............................................................ 21
The Origin of the Arachnids: A Palaeofantasy .......................................... 22
Arachnids and the Colonisation of Land .................................................. 25
Fossil Mites ............................................................................................... 26
Fossil Acariformes ............................................................................... 27
Fossil Parasitiformes ............................................................................ 27
Potential Arachnid Relatives of Mites ...................................................... 28
Palpigradi ............................................................................................. 29
Opiliones .............................................................................................. 31
Ricinulei ............................................................................................... 32
Pseudoscorpionida. ............................................................................... 33
Solifugida ............................................................................................. 34
Summary and Preview .............................................................................. 34
References ................................................................................................. 35
3 Systematic and Morphological Survey .................................................. 39
What Is ‘Acari’? The Question of Mite Monophyly ................................. 39
Parasitiformes: Ticks and Their Relatives ................................................. 40
Acariformes: The Mite-Like Mites ........................................................... 50
ix