Proc. R

. Soc. B

| vol. 277 n

o. 1690 p

p. 1937–2095 | 7 July 2010

7 July 2010

volume 277 . number 1690 . pages 1937–2095

ISSN 0962-8452

volume 277

number 1690

pages 1937–2095

7 July 2010

Founded in 1660, the Royal Society is the independent scientific academy of the UK, dedicated to promotingexcellence in science

Registered Charity No 207043rspb.royalsocietypublishing.orgPublished in Great Britain by the Royal Society, 6–9 Carlton House Terrace, London SW1Y 5AG

Review articlesAtmospheric oxygen level and the evolution of insect body size 1937J. F. Harrison, A. Kaiser & J. M. VandenBrooks

Research articlesPartner choice in Medicago Truncatula–Sinorhizobium symbiosis 1947C. Gubry-Rangin, M. Garcia & G. Béna

Ergatoid queen development in the ant Myrmecina nipponica: modular and heterochronic regulation of caste differentiation 1953S. Miyazaki, T. Murakami, T. Kubo, N. Azuma, S. Higashi & T. Miura

The evolution of phototransduction from an ancestral cyclic nucleotide gated pathway 1963D. C. Plachetzki, C. R. Fong & T. H. Oakley

The shrinking ark: patterns of large mammal extinctions in India 1971K. K. Karanth, J. D. Nichols, K. U. Karanth, J. E. Hines & N. L. Christensen Jr

How fast could HIV change gene frequencies in the human population? 1981D. Cromer, S. M. Wolinsky & A. R. McLean

Fossil avian eggshell preserves ancient DNA 1991C. L. Oskam, J. Haile, E. McLay, P. Rigby, M. E. Allentoft, M. E. Olsen, C. Bengtsson, G. H. Miller, J.-L. Schwenninger, C. Jacomb, R. Walter, A. Baynes, J. Dortch, M. Parker-Pearson, M. T. P. Gilbert, R. N. Holdaway, E. Willerslev & M. Bunce

MHC gene copy number variation in Tasmanian devils: implications for the spread of a contagious cancer 2001H. V. Siddle, J. Marzec, Y. Cheng, M. Jones & K. Belov

Selfish strategies and honest signalling: reproductive conflicts in ant queen associations 2007L. Holman, S. Dreier & P. d’Ettorre

Acellular pertussis vaccination facilitates Bordetella parapertussis infection in a rodent model of bordetellosis 2017G. H. Long, A. T. Karanikas, E. T. Harvill, A. F. Read & P. J. Hudson

The asymmetry of the carpal joint and the evolution of wing folding in maniraptoran theropod dinosaurs 2027C. Sullivan, D. W. E. Hone, X. Xu & F. Zhang

Reproductive competition promotes the evolution of female weaponry 2035N. L. Watson & L. W. Simmons

Dynamics and ecological consequences of avian influenza virus infection in greater white-fronted geese in their winter staging areas 2041D. Kleijn, V. J. Munster, B. S. Ebbinge, D. A. Jonkers, G. J. D. M. Müskens, Y. Van Randen & R. A. M. Fouchier

A naturally occurring variant of Hsp90 that is associated with decanalization 2049C. M. Sgrò, B. Wegener & A. A. Hoffmann

Monitoring long-term evolutionary changes following Wolbachia introduction into a novel host: the Wolbachia popcorn infection in Drosophila simulans 2059L. B. Carrington, A. A. Hoffmann & A. R. Weeks

A developmental perspective on the evolution of sexual size dimorphism of a moth 2069R. C. Stillwell & G. Davidowitz

Dispersal capacity and diet breadth modify the response of wild bees to habitat loss 2075R. Bommarco, J. C. Biesmeijer, B. Meyer, S. G. Potts, J. Pöyry, S. P. M. Roberts, I. Steffan-Dewenter & E. Öckinger

An immunological cost of begging in house sparrow nestlings 2083G. Moreno-Rueda

Interactions among social monitoring, anti-predator vigilance and group size in eastern grey kangaroos 2089F.-R. Favreau, A. W. Goldizen & O. Pays

See further with the Royal Society in 2010 – celebrate 350 years

RSPB_277_1690_Cover.qxd 05/12/10 09:58 AM Page 1

Publishing Editor: Victoria Millen

(tel: +44 (0)20 7451 2631; fax: +44 (0)20 7976 1837;

victoria.millen@royalsociety.org)

Editorial Coordinator: Charlotte Wray

(tel: +44 (0)20 7451 2629; proceedingsb@royalsociety.org)

Senior Production Editor: Siân Roderick

6–9 Carlton House Terrace, London SW1Y 5AG, UK

rspb.royalsocietypublishing.org

AIMS AND SCOPE Research papers. Proc. R. Soc. B welcomes articles of high quality in all aspects ofbiology. Currently, it is particularly strong in ecology in the widest sense, and behaviouraland evolutionary biology. Proc. R. Soc. B has also published significant numbers of papersin epidemiology, human biology, neuroscience, palaeontology and biomechanics. Authorsare encouraged to submit articles not only in these fields, but in all branches of biology.The criteria for selection are scientific excellence, originality and potential interest to awide spectrum of biologists.Review papers. The journal also publishes topical reviews in emerging areas of biologicalscience that will be of interest to non-specialists. Topical reviews are commissioned butunsolicited submissions will be considered.Comment papers. Short additions to, or criticisms of, papers already published in thisjournal will also be considered. At the discretion of the editors, the authors of the originalpaper may be invited to reply. Comment papers should be short, interesting and conformto the rules of good manners.

Editorial Board

M. P. Hassell, Editor-in-Chief, Department of Life Sciences, Imperial College LondonE. Adkins-Regan, Editor, Department of Psychology,Cornell UniversityN. C. Bennett, Editor, Department of Zoology andEntomology, University of PretoriaI. C. Cuthill, Editor, School of Biological Sciences,University of BristolJ. A. P. Heesterbeek, Editor, Department of FarmAnimal Health, University of UtrechtM. T. Siva-Jothy, Reviews Editor, Department ofAnimal & Plant Sciences, University of Sheffield

F. W. Allendorf, Division of Biological Sciences,University of MontanaS. H. Alonzo, Department of Ecology and EvolutionaryBiology, Yale UniversityA. T. D. Bennett, School of Life and EnvironmentalSciences, Deakin UniversityM. W. Blows, Evolutionary Quantitative GeneticsGroup, University of QueenslandM. Bonsall, Department of Zoology, University of OxfordR. Boyd, Department of Anthropology, University ofCalifornia, Los AngelesP. M. Brakefield, Institute of Zoology, Leiden Universityand Department of Animal and Plant Sciences,University of SheffieldR. Bshary, Institute of Zoology, University of NeuchâtelN. Burgess, Institute of Cognitive Neuroscience andDepartment of Anatomy, University College LondonM. Chapuisat, Department of Ecology and Evolution,University of LausanneL. Chittka, School of Biological and Chemical Sciences,Queen Mary, University of LondonS. L. Chown, Department of Botany and Zoology,University of StellenboschR. L. Cifelli, Department of Zoology and School ofGeology and Geophysics, University of OklahomaA. S. Clare, Marine Science and Technology, Universityof Newcastle upon TyneN. S. Clayton, Department of ExperimentalPsychology, University of CambridgeA. Cockburn, School of Biology, Australian NationalUniversityD. P. Costa, Center for Ocean Health University of California, Santa CruzG. Cowlishaw, Institute of Zoology, University of Cambridge

S. R. X. Dall, School of Biosciences, University of Exeterat CornwallM. S. Dawkins, Department of Zoology, University of OxfordT. Day, Departments of Mathematics & Biology,Queen's UniversityL. De Meester, Laboratory of Aquatic Ecology,Katholieke Universiteit LeuvenG. Edgecombe, Palaentology Department, Natural History MuseumS. F. Elena, Instituto de Biología Molecular y Celular dePlantas (IBMCP), CSIC-UPVJ. P. Evans, School of Animal Biology, The University ofWestern AustraliaC. G. Faulkes, School of Biological Sciences, QueenMary, University of LondonR. G. Foster, Department of Ophthalmology,University of OxfordT. Fukatsu, National Institute of Advanced IndustrialScience and Technology (AIST), JapanD. M. Gordon, Department of Biology, StanfordUniversityR. D. Gray, Psychology Department, University of AucklandL. D. Harder, Department of Biological Sciences,University of CalgaryR. G. Harrison, Department of Ecology andEvolutionary Biology, Cornell UniversityS. Healy, School of Psychology, University of St AndrewsJ. A. Hutchings, Department of Biology, Life ScienceCentre, Dalhousie UniversityM. Kirkpatrick, College of Natural Sciences, Universityof Texas at AustinR. Knapp, Department of Zoology, University ofOklahomaJ. C. Koella, Faculty of Life Sciences, Imperial CollegeLondonS. P. Long, Department of Plant Biology, University of IllinoisI. J. Lovette, Cornell Laboratory of Ornithology,Cornell UniversityB. S. Low, School of Natural Resources andEnvironment, University of MichiganP. Lundberg, Department of Theoretical Ecology, Lund UniversityJ. R. Madden, School of Psychology, University of ExeterL. B. Martin, Department of Integrative Biology,University of South Florida

A. E. Minetti, Department of Human Physiology,University of MilanM. Noor, Biology Department, Duke UniversityK. Norris, School of Agriculture, Policy andDevelopment, University of ReadingD. H. Nussey, School of Biological Sciences, Universityof EdinburghT. Pizzari, Department of Zoology, University of OxfordT. D. Price, Department of Ecology and Evolution, The University of ChicagoD. Promislow, Department of Genetics, University of GeorgiaA. Purvis, Faculty of Natural Sciences, Biology Division,Imperial College LondonL. Råberg, Department of Animal Ecology, LundUniversityA. N. Radford, School of Biological Sciences, Universityof BristolJ. Radwan, Institute of Environmental Sciences,Jagiellonian UniversityM. Rees, Department of Animal and Plant Sciences,University of SheffieldS. S. Renner, Department of Biology, University of MissouriL. Rowe, Department of Ecology & EvolutionaryBiology, University of Toronto H. D. Rundle, Department of Biology, University ofOttawaD. E. Ruzzante, Department of Biology, DalhousieUniversityW. Schultz, Department of Physiology, Development and Neuroscience, University ofCambridgeL. W. Simmons, Centre for Evolutionary Biology,University of Western AustraliaJ. E. Smith, Center for Marine Biodiversity andConservation, Scripps Institution of OceanographyM. J. Telford, Department of Genetics, Evolution andEnvironment, University College LondonH. Tsukaya, Graduate School of Science, University of TokyoJ. L. van Leeuwen, Experimental Zoology Group,Wageningen UniversityJ. T. Wootton, Department of Ecology & Evolution, The University of ChicagoXing Xu, Institute of Vertebrate Paleontology andPaleoanthropology, BeijingDa-yong Zhang, Beijing Normal University

Copyright © 2010 The Royal SocietyExcept as otherwise permitted under the Copyright, Designs and Patents Act, 1988, this publication may only be reproduced, stored or transmitted, in any form orby any other means, with the prior permission in writing of the publisher, or in the case of reprographic reproduction, in accordance with the terms of a licenceissued by the Copyright Licensing Agency. In particular, the Society permits the making of a single photocopy of an article from this issue (under Sections 29 and38 of this Act) for an individual for the purposes of research or private study.

Typeset in India by Techset Composition Limited, Salisbury, UK. Printed by Latimer Trend, Plymouth.

This paper meets the requirements of ISO 9706:1994(E) and ANSI/NISO Z39.48-1992 (Permanence of Paper) effective with volume 247, issue 1318, 1992.

Proceedings of the Royal Society B (ISSN 0962-8452) is published twice a month for $2128 per year by the Royal Society, and is distributed in the USA by Agent named AirBusiness, C/O Worldnet Shipping USA Inc., 149-35 177th Street, Jamaica, New York, NY11434, USA. US Postmaster: Send address changes to Proceedings of the Royal Society B,C/O Air Business Ltd, C/O Worldnet Shipping USA Inc., 149-35 177th Street, Jamaica, New York, NY11414.

Editor-in-ChiefProfessor Michael P. HassellReviews EditorProfessor Michael T. Siva-JothyPublishing EditorVictoria Millen

Editors:Professor Elizabeth Adkins-ReganProfessor Nigel BennettProfessor Innes CuthillProfessor Hans Heesterbeek

Subscription prices All other2010 calendar year Europe USA & Canada countries

Printed version plus electronic £1106/€1437 US$2128 £1231/US$2207access

SUBSCRIPTIONSIn 2010 Proc. R. Soc. B (ISSN 0962-8452) will be published twice a month.Full details on how to subscribe can be obtained either by contacting ourjournal fulfilment agent, Portland Customer Services, Commerce Way,Colchester CO2 8HP. Tel: +44 (0)1206 796351; fax: +44 (0)1206 799331;email: sales@portland-services.com or by visiting our website atroyalsocietypublishing.org/info/subscriptions. The Royal Society is aRegistered Charity No. 207043.

The Royal Society, the national academy of science ofthe UK and the Commonwealth, is at the cutting edgeof scientific progress. We support many top youngscientists, engineers and technologists, influencescience policy, debate scientific issues with the publicand much more. We are an independent, charitablebody and derive our authoritative status from over 1400Fellows and Foreign Members.

During 2010, we are celebrating the Royal Society’s350th anniversary. As part of this, there will be anexciting programme of activities – exhibitions, lectures,conferences, a new book, a vast science festival on theSouth Bank in London, television and radiobroadcasting and much more besides.

Our mission: to expand knowledge and further the roleof science and engineering in making the world a betterplace.

The Royal Society’s strategic priorities are to:

• invest in future scientific leaders and ininnovation,

• influence policymaking with the best scientificadvice,

• invigorate science and mathematics education,• increase access to the best science internationally,

and• inspire an interest in the joy, wonder and

excitement of scientific discovery.

For further information on the Society’s activities, pleasecontact the following departments on the extensionslisted by dialling +44 (0)20 7839 5561, or visit theSociety’s Web site (royalsociety.org).

Research Support (UK grants and fellowships)Research Appointments (Fellowships): 2542Research Grants: 2223International travel Grants: 2555Newton International Fellowships: 2559

Science AdviceScience Policy Centre: 2550

Science CommunicationGeneral enquiries: 2573

Library and Information ServicesLibrary/archive enquiries: 2606

Selection criteria

The criteria for selection are scientific excellence, originality and

interest across disciplines within biology. The Editors are responsible

for all editorial decisions and they make these decisions based on

the reports received from the referees and/or Editorial Board

members.

Many more good manuscripts are submitted to us than we have

space to print, and we give preference to those that present

significant advances of broad interest. Submission of preliminary

reports, of articles that merely confirm previous findings, and of

articles that are likely to interest only small groups of specialists, is

not encouraged. All articles are sent to Editorial Board members for

an initial assessment of their suitability, and may be returned to

authors without in-depth peer review if this assessment makes it

seem unlikely that they will be accepted.

Publishing format

Proc. R. Soc. B articles are published regularly online and in print

issues twice a month. Along with all Royal Society journals, we are

committed to archiving and providing perpetual access. Although

papers are limited to ten journal pages in length (with a per-page

charge for articles exceeding six pages), there is the facility for

including Electronic Supplementary Material (ESM). Contents of the

ESM might include details of methods, derivations of equations,

large tables of data, DNA sequences and computer programs.

However, the printed version must include enough detail to satisfy

most non-specialist readers. Supplementary data up to 10Mb is

placed on the Society's website free of charge. Larger datasets must

be deposited in recognised public domain databases by the author.

Conditions of publication

Articles must not have been published previously, nor be under

consideration for publication elsewhere. The main findings of the

article should not have been reported in the mass media. Like many

journals, Proc. R. Soc. B employs a strict embargo policy where the

reporting of a scientific article by the media is embargoed until a

specific time. The Editor-in-Chief has final authority in all matters

relating to publication.

Electronic Submission details

The Royal Society’s electronic-submission and peer-review service

provides Proc. R. Soc. B authors with the facility to submit their

papers online. The service allows you to upload files in a reliable and

user-friendly way, using a Web-based system. When your paper is

received, an immediate acknowledgement is sent that details how

you can track your contribution online.

For full submission guidelines and access to all journal content please

visit the Proc. R. Soc. B website, for further details see

rspb.royalsocietypublishing.org.

Cover image: The head region of the cnidarian Hydra magnipapillata as revealed by confocal microscopy. Neurons are shown in red,

musculature is shown in green and cell nuclei are shown in blue. The mouth opening of the animal is at the centre of the image.

(See pages 1963–1969; image courtesy of David C. Plachetzki.)

GUIDANCE FOR AUTHORS

RSPB_277_1690_Cover.qxd 05/12/10 09:58 AM Page 2