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First published in 1963, Advances in Parasitology contains comprehensive and up-to-date reviews in all areas of interest in contemporary parasitology. Advances in Parasitology includes medical studies on parasites of major influence, such as Plasmodium falciparum and Trypanosomes. The series also contains reviews of more traditional areas, such as zoology, taxonomy, and life history, which shape current thinking and applications. With an impact factor of 3.9 the series ranks second in the ISI Parasitology subject category.

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Advances in PARASITOLOGY VOLUME 60 i Editorial Board M. Coluzzi, Director, Istituto de Parassitologia, Universita` Degli Studi di Roma ‘La Sapienza’, P. le A. Moro 5, 00185 Roma, Italy C. Combes, Laboratoire de Biologie Animale, Universite´ de Perpignan, Centre de Biologie et d’Ecologie Tropicale et Me´diterrane´enne, Avenue de Villeneuve, 66860 Perpignan Cedex, France D.D. Despommier, Division of Tropical Medicine and Environmental Sciences, Department of Microbiology, Columbia University, 630 West 168th Street, New York, NY 10032, USA J.J. Shaw, Instituto de Cieˆncias Biome´dicas, Universidade de Sa˜o Paulo, av. Prof. Lineu Prestes 1374, 05508-900, Cidade Universita´ria, Sa˜o Paulo, SP, Brazil K. Tanabe, Laboratory of Biology, Osaka Institute of Technology, 5-16-1 Ohmiya Asahi-Ku, Osaka 535, Japan ii Advances in PARASITOLOGY Edited by J.R. BAKER Royal Society of Tropical Medicine and Hygiene, London, England R. MULLER London School of Hygiene and Tropical Medicine, London, England and D. ROLLINSON The Natural History Museum, London, England VOLUME 60 ELSEVIER ACADEMIC PRESS Amsterdam Boston Heidelberg London New York Oxford Paris San Diego San Francisco Singapore Sydney Tokyo iii ELSEVIER B.V. Radarweg 29 P.O. Box 211 1000 AE Amsterdam The Netherlands ELSEVIER Ltd The Boulevard, Langford Lane, Kidlington Oxford OX5 1GB UK ELSEVIER Inc. 525 B Street, Suite 1900 San Diego CA 92101-4495 USA ELSEVIER Ltd 84 Theobalds Road London WC1X 8RR 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, inc