The biology of centipedes

483 Pages·1981·7.894 MB·English

by  Lewis, J.G.E.

The biology of centipedes

J.G.E.LEWIS
The biology of centipedes
CAMBRIDGE UNIVERSITY PRESS
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© Cambridge University Press 1981
This publication is in copyright. Subject to statutory exception
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no reproduction of any part may take place without
the written permission of Cambridge University Press.
First published 1981
This digitally printed first paperback version 2006
A catalogue record for this publication is available from the British Library
ISBN-13 978-0-521-23413-9 hardback
ISBN-10 0-521-23413-1 hardback
ISBN-13 978-0-521-03411-1 paperback
ISBN-10 0-521-03411-6 paperback
CONTENTS
Preface vii
1 Introduction 1
2 External morphology and functional anatomy 7
3 The integument, moulting and regeneration 47
4 The musculature and endoskeleton 67
5 The nervous system and sense organs 85
6 Sensory responses and related behaviour 119
7 Endocrinology 131
8 The alimentary canal 141
9 The poison glands 156
10 Feeding and digestion 167
11 The respiratory system 189
12 The circulatory system 209
13 Pigments 220
14 Connective tissue and fat body 226
15 Head glands 230
16 The Malpighian tubules and nephridia 239
17 The reproductive system and reproduction 252
18 Post-embryonic development and life history 299
19 Epidermal glands and their function, defence and predators 337
20 Parasites 359
21 Physiology and ecology 375
22 Taxonomy 407
23 Relationships of the chilopod orders 418
24 The classification of the Chilopoda 425
Bibliography 429
Index 457

PREFACE
This book is an attempt to bring together information on centi-
pedes hitherto to be found only in old texts or scattered through
the literature and in unpublished theses.
My interest in the group is due to Sir Eric Smith, FRS who, as
Professor J. E. Smith, suggested that I carry out postgraduate
research on littoral centipedes. Since that time I have had a great
deal of advice and encouragement from Mr. J. Gordon Blower, Dr
E. H. Eason, Dr R. E. Crabill, Jr and Professor J. L. Cloudsley-
Thompson. Over the years, I have received much help from mem-
bers of the Arachnid Section of the British Museum (Natural
History) most notably, Mr K. H. Hyatt, Mr D. Macfarlane, Mr F.
R. Wanless and the late Mr D. J. Clark and also from the library staff
of the Museum.
The Headmaster and Governors of Dover College generously
allowed me a term's leave of absence to take up a School
Teacher Fellow-Commonership at Sidney Sussex College, Cambridge,
in order to complete the book. I am greatly indebted to
the Master and Fellows of Sidney Sussex for their help and
hospitality and to the librarians of the Cambridge Zoology
Department Library for their assistance. I am grateful to Dr D. L.
Gunn, CBE and the Royal Sbciety Research in Schools Committee for
their support during the preparation of this book.
I am likewise indebted to Miss K. E. Yule for help with French
and German and to Mrs K. Stefan for translating the Russian
papers. My thanks are also due to Mrs B. Bradford for typing the
manuscript and to the staff of Cambridge University Press for their
advice during its processing for publication.
Above all, my thanks are due to my wife, for checking the manu-
script and for constant advice, support and encouragement.
to S.J.C.
Introduction
Centipedes (members of the arthropodan class Chilopoda) are
common and relatively familiar animals which are found in soil and
litter or under stones or bark. They are soft-bodied and dorso-
ventrally flattened having from 15 to 181 pairs of legs, one pair to
each trunk segment. Species from temperate regions are usually of
moderate size, varying from 1 to 10 cm in length and of drab
brownish or yellowish coloration. Many tropical species of the
order Scolopendromorpha are large, one reaches a length of 26 cm,
and are brightly coloured: red, black and orange, green or violet.
Like other arthropods, the centipedes are bilaterally symmetrical,
metamerically segmented animals with a double ventral nerve cord,
typically with a ganglion in each segment and concentrations of
nervous tissue above and below the gut at the anterior end of the
body. A circulatory system is present carrying blood forwards in a
dorsal vessel and backwards in a ventral vessel. The body is
covered by a non-living layer, the cuticle, which is secreted by the
epidermis. The cuticle is in the form of relatively rigid sclerites
separated by flexible arthrodial membranes; it is shed periodically
to allow growth, a phenomenon known as moulting or ecdysis. The
anterior part of the body is differentiated to form a head which
bears a pair of antennae, a pair of jaws (mandibles) and two pairs
of jointed legs modified to form mouthparts (the first and second
maxillae). The legs of the first trunk segment are modified to form
the characteristic poison claws which are used to seize prey.
The centipedes have often been grouped as the Myriapoda with
the millipedes (Class Diplopoda) and two classes of soil-dwelling
arthropods: the Symphyla with 12 pairs of legs and a pair of stout
anal cerci and the minute Pauropoda with nine or ten pairs of legs
and biramous antennae. Myriapods may be defined as arthropods
with one pair of antennae, two or three pairs of mouthparts and
numerous pairs of legs.
Introduction 2
Opinions as to the status of the four groups of myriapods have
varied considerably. They have sometimes been regarded as four
classes showing little true relationship to each other. Manton
(1970), however, has discussed the reasons for the reinstatement of
the Myriapoda as a natural group. A study of the jaw and leg
Fig. 1. Geophilus carpophagus, body length 40 mm (after a
photograph by Manton in Eason, 1964).