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Advances in Insect Physiology publishes eclectic volumes containing important, comprehensive and in-depth reviews on all aspects of insect physiology. It is an essential reference source for invertebrate physiologists and neurobiologists, entomologists, zoologists and insect biochemists. First published in 1963, the serial is now edited by Steve Simpson (Oxford University, UK).

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Advances in Insect Physiology Volume 32 i This page intentionally left blank ii Advances in Insect Physiology edited by S. J. Simpson School of Biological Sciences, The University of Sydney, Sydney, Australia Volume 32 ELSEVIER ACADEMIC PRESS Amsterdam  Boston  Heidelberg  London  New York  Oxford  Paris  San Diego  San Francisco  Singapore  Sydney  Tokyo iii ELSEVIER Ltd The Boulevard, Langford Lane Kidlington, Oxford OX5 1GB, UK r 2005 Elsevier Ltd. All rights reserved. This work is protected under copyright by Elsevier Ltd, and the following terms and conditions apply to its use: Photocopying Single photocopies of single chapters may be made for personal use as allowed by national copyright laws. Permission of the Publisher and payment of a fee is required for all other photocopying, including multiple or systematic copying, copying for advertising or promotional purposes, resale, and all forms of document delivery. Special rates are available for educational institutions that wish to make photocopies for non-profit educational classroom use. Permissions may be sought directly from Elsevier’s Rights Department in Oxford, UK: phone (+44) 1865 843830, fax (+44) 1865 853333, e-mail: [email protected] Requests may also be completed on-line via the Elsevier homepage (http://www.elsevier.com/locate/permissions). In the USA, users may clear permissions and make payments through the Copyright Clearance Center, Inc., 222 Rosewood Drive, Danvers, MA 01923, USA; phone: (+1) (978) 7508400, fax: (+1) (978) 7504744, and in the UK through the Copyright Licensing Agency Rapid Clearance Service (CLARCS), 90 Tottenham Court Road, London W1P 0LP, UK; phone: (+44) 20 7631 5555; fax: (+44) 20 7631 5500. Other countries may have a local reprographic rights agency for payments. Derivative Works Tables of contents may be reproduced for internal circulation, but permission of the Publisher is required for external resale or distribution of such material. Permission of the Publisher is required for all other derivative works, including compilations and translations. Electronic Storage or Usage Permission of the Publisher is required to store or use electronically any material contained in this work, including any chapter or part of a chapter. Except as outlined above, no part of this work may be reproduced, stored in a retrieval system or transmitted in any form or by any means, electronic, mechanical, photocopying, recording or otherwise, without prior written permission of the Publisher. Address permissions requests to: Elsevier’s Rights Department, at the fax and e-mail addresses noted above. Notice No responsibility is assumed by the Publisher for any injury and/or damage to persons or property as a matter of products liability, negligence or otherwise, or from any use or operation of any methods, products, instructions or ideas contained in the material herein. Because of rapid advances in the medical sciences, in particular, independent verification of diagnoses and drug dosages should be made. First edition 2005 British Library Cataloguing in Publication Data A catalogue record is available from the British Library. Academic Press An imprint of Elsevier 525 B Street, Suite 1900 San Diego, CA 92101-4495, USA http://www.academicpress.com
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