CINXE.COM

<!DOCTYPE html><html lang="en-US"><head><title>Just a moment...</title><meta http-equiv="Content-Type" content="text/html; charset=UTF-8"><meta http-equiv="X-UA-Compatible" content="IE=Edge"><meta name="robots" content="noindex,nofollow"><meta name="viewport" content="width=device-width,initial-scale=1"><style>*{box-sizing:border-box;margin:0;padding:0}html{line-height:1.15;-webkit-text-size-adjust:100%;color:#313131;font-family:system-ui,-apple-system,BlinkMacSystemFont,Segoe UI,Roboto,Helvetica Neue,Arial,Noto Sans,sans-serif,Apple Color Emoji,Segoe UI Emoji,Segoe UI Symbol,Noto Color Emoji}body{display:flex;flex-direction:column;height:100vh;min-height:100vh}.main-content{margin:8rem auto;max-width:60rem;padding-left:1.5rem}@media (width <= 720px){.main-content{margin-top:4rem}}.h2{font-size:1.5rem;font-weight:500;line-height:2.25rem}@media (width <= 720px){.h2{font-size:1.25rem;line-height:1.5rem}}#challenge-error-text{background-image:url(data:image/svg+xml;base64,PHN2ZyB4bWxucz0iaHR0cDovL3d3dy53My5vcmcvMjAwMC9zdmciIHdpZHRoPSIzMiIgaGVpZ2h0PSIzMiIgZmlsbD0ibm9uZSI+PHBhdGggZmlsbD0iI0IyMEYwMyIgZD0iTTE2IDNhMTMgMTMgMCAxIDAgMTMgMTNBMTMuMDE1IDEzLjAxNSAwIDAgMCAxNiAzbTAgMjRhMTEgMTEgMCAxIDEgMTEtMTEgMTEuMDEgMTEuMDEgMCAwIDEtMTEgMTEiLz48cGF0aCBmaWxsPSIjQjIwRjAzIiBkPSJNMTcuMDM4IDE4LjYxNUgxNC44N0wxNC41NjMgOS41aDIuNzgzem0tMS4wODQgMS40MjdxLjY2IDAgMS4wNTcuMzg4LjQwNy4zODkuNDA3Ljk5NCAwIC41OTYtLjQwNy45ODQtLjM5Ny4zOS0xLjA1Ny4zODktLjY1IDAtMS4wNTYtLjM4OS0uMzk4LS4zODktLjM5OC0uOTg0IDAtLjU5Ny4zOTgtLjk4NS40MDYtLjM5NyAxLjA1Ni0uMzk3Ii8+PC9zdmc+);background-repeat:no-repeat;background-size:contain;padding-left:34px}@media (prefers-color-scheme:dark){body{background-color:#222;color:#d9d9d9}}</style><meta http-equiv="refresh" content="390"></head><body class="no-js"><div class="main-wrapper" role="main"><div class="main-content"><noscript><div class="h2"><span id="challenge-error-text">Enable JavaScript and cookies to continue</span></div></noscript></div></div><script>(function(){window._cf_chl_opt={cvId: '3',cZone: "royalsocietypublishing.org",cType: 'managed',cRay: '8e9d37886eee9c5d',cH: '41RLshhpYW7CxXaME82allLj.BbsQ3Xd88D6Qga0FSE-1732826886-1.2.1.1-yawSWDmhqNUVa_EWAFqxkGnSSgMGKAjMoi.6FrGbRY05GUZtAPM9AMxPgGz60JxG',cUPMDTk: "\/doi\/10.1098\/rsbl.2014.0911?__cf_chl_tk=FcKSCv99bfgPY.Za89vGIy05tqG5guaphRU17pHZfKM-1732826886-1.0.1.1-U4AxeuiRHQn27LFznTKugh85disMZF4l4qTzEJhFTEE",cFPWv: 'b',cITimeS: '1732826886',cTTimeMs: '1000',cMTimeMs: '390000',cTplC: 0,cTplV: 5,cTplB: 'cf',cK: "",fa: "\/doi\/10.1098\/rsbl.2014.0911?__cf_chl_f_tk=FcKSCv99bfgPY.Za89vGIy05tqG5guaphRU17pHZfKM-1732826886-1.0.1.1-U4AxeuiRHQn27LFznTKugh85disMZF4l4qTzEJhFTEE",md: "ezA2o5ATTaZjHLTKi6rbnrRdF8i13hQGMOePhzeWz4E-1732826886-1.2.1.1-Lut_82hV2NLWW85bScRraTdkCzSJEUgQQ7fgEQ_.qnqpAzL1JhnTbbAI5_.y5Emj9LjxaPkLMF2fPRvqMqXEbFLXfmKQFWwVy.BaSA0YQZX.jxODRSAIhpeCCw9x8izBGEmM.K3nRrn6J261TYFKMJddJWuORK7hLUVF.FuJDj2afL_dFcaZJMsSImCdo7D0lH6qNbvw.g1fFFXizvaEv2jsFMHbPX.rkpqUcAxMwuWp2NrHa5HAxX1qxx245mY.Dtuf_4J7tBJcctzrBXHIuuPNZBV_aAvjKRTvrkbM4r5g1AFFGgNOnq5w8g690INpN3iZ3qgTRTcy6IsxvtM6KniDtDdTzNZAXAGZJOBeKSykvEXS8K6Z2krzGnsaeyaP12CHT3NeUkK6XmFV6ttKtyNLrEd79_KhmT9QDeJjZAxrw0Uaf3YOzELrmMwIPZ3QM5FTq1GUdtj12GxleRtJU.Ikc4iJSMrPv_7RA4yoP8ALLDuJNSVL8vZFRRqcaunlo9.EFcyXKGzDdh_yjUYHoiSYjGgKrmVGc6h.MnEpDpJTP7jwGOVI0_zmxEeEuJk_.UGf_eWQjQF6iXk0L6Myl3G2LPtHHgCkf5VczQWI.RR4wR4Q31o73nmwPfmKKneiKWZ3Eettdz7E4CqpYa6RbQ7LupI6sJpDeNx3Xe_I2MzcziZLhFhCtZM0b.N20IkD.C___OcFsM0GYka_6I33zSWqtrsabZK7O0kIZUejC7Bgufsniwna3KAbXeUU1WapPkmFjO8XAT5Y5ZxHuHll7QkdXmRLKLYwXK7J4HOOH.zjAK5WzlRG_AgQUG52mcYaxxwhenyeQYQYPLzgCPwmfEKm3gNS5TRV3NwHo5JfJ_p74WdZ79SFXDeovZebKzFmD4BOM_TaVnXSDSLrsiVwyGRIZTpg50JL6OVY6zMseNDRCin9DonUyK2R9lK.J6gc7dT4gTgxbWCvVo46PkHPGd5lvh7rihZSuMR3.zVk3BoKKKV1DLXpKpCuPChvBFWeqnbPknn3Nu2_iVdKcoTHjE5do0cTnIhuQG9KE__HqTkxIeSzEJB4AUaOoEFUqdRARrKXQCYfaNG6HAPPGn578yjJsRnXRvvhVdRQNDeTa9mCatxZXp0qbsB_7RAChxPsCpXI120PAurjuOx8OcQ827KkYnCXLsNCyR5biMBTY442k0QxCCZezuosqapMkqULUJLvUtuUgkjUh67Z.ADqrWoAQ_1xed787oQ2CR6zNf22eBv6NSwHPYEgKeB2ntR7e6_GxC9sPRRAZ12uq1rJCgqxf27LzTkhLB9GlS2iCadw5RBfsu8oNwwGq3t92vrjSL.7UlhSR3WPxzLgtpCuh1tRS8NqwKzwbBxU1CGyF6dm.de6H9VaOoO_JYel76.nkuhfo6Z3EILsik.wdg800B_OzK25hbHIBFReVPlC7a4A9n10MznhMuaOFzjiC1jwsbXm.pSQLp3_UcYtx7Evd0kojwQQ.0cmuLxJdDe_okqsfBiOzy8cS2gFFsi9ANlTClsPmI7EyLCKRwSo6KOL9PkFmWZoKhH1trlfE8EFEudlq.lk7GkVZhVYb9LWtYjk8mnjXuryMx0L2S066VQsIV3uiLM921WbBDyg6CAy8Fx2mUHY_0ENTK0GknRw7wJ7RbduLU7LNbjhGrFPpLbQSqoW5VZA3Z87YxgIUXgqckj1.ot3E80glkKDidNn0MzsNAh52_8FVA74a2PioBJZSYWN8iOXv2AjigiogAxw9Z3678c0cOknHDyGPqbIQYEAP8AvDoI3hl0pn6vgyujk4H6tU9JhhtxzHkCEd70BpCsdHFpjM4iQ1o1Hb2GdN_Akzf31YvGAeNnFy3kEXtF56Lly9RMfpPXQokjeJOF2kVU8HOkraxlljEuddAE.XsFgF8OAa4OE.6K0iihF_fYmZ3x3JhJAqh0xw0a8X0AKJ_Jj3oMYn7ASPu86AC8mL0Kk1q2uyvWcPwerRmsRMJjWmVr4KEdjzDxOY2Xyz2bDwNHgKaBIvcceV_lJXFfFZ4tVOR1eXJDPChJR56I9OfX6baexdNf2mRPP73qo.9_y1xPZ3HODUCmoLJQ1mOjkWmWrlfbSdsPf1JqlZ_B9x1FwDjxJZlWQcDJf7Rf9MaWd.cmr4hUVIfpC13htCmx62ZtHJtCJNBVlgaVSeJxyxwfnjEh_vXNCtXn12Ocbb9ppLJN1OZizB23CTkZSlz..ykoWww88XL8cxps1GYxAqK68WYyymp.R7yBY2SqFMvbIbfFj56j5c8jXT2VKmFc0kA_uFvLX4g3E9nKSJrWqhh2siJ.p971NUMvDngIGII5vNyn1bmr_cX.ALriRskKrKRQFfKqEajPFX5IS09YtqSSJjRFIp.Jp.WHEqis1CIX6x_DgAihYuTZmbodTqHjeFloZ2ZWFyLSK7dV1psmRlRgagXTJHLGDxFjciTtI8rYVAGz40arqRUJ6p9Hxvw6b1Cm9KUbCo9UU3Pqr.Q.oH16vMgnUV12fCNXjj7Xyx8KNNHknmx2NLkwEYcx4vrVHWNnYx6zkbk9J4IRe8NkRt3If0noAsEWJ9daQ2281KdhyF9wbvp28GBRy7FTXLAX5fsdyLJZmGVV26.yd8NWAQBoUhjOQrxoe.Fng0zDg_hxiBPBiLmDjzZ4u_SVy_J2pNPwA9d1QSPS3Tv2rNWWTWZjDuPcSIGXxrrtpht.W8XyZvxt0thcosrqWSEaNZP.jKOKREbzMt9q77iDvov_FwoV1lPnFyWzrfpEFIelcdJ_w1RWN4hjUjgXymwrp7o9Ltwlkaa1WhbDNKE6_FqSpuNeX16hTCZz2bWO4qB5NTjRW9UFkr3FbE94OAHHkrls1ApOzQQh7TrKnJ_E0mlsgJg1RKCUy2i3zEmISHxY2hn.nBfOzlJ7clxNlh2nYMA6ZQqsEqslwz.RrIqSddaOufDVZ65W3jPl0rw4FwXz2s_yIumr79dq.ad6XLAqwgyv3Q_fD",mdrd: "eoN1DbwV8CLpbZ32U_Gc1LEJkjwH4FDhsmbd9MGeTQw-1732826886-1.2.1.1-o9Q.RozS3oz.aUGWspsJtXfq9X2X0scbBL36UKQRDxO.GTYdOqXn68My7dUVARTt.Y0ytq6olWCwRMz5pGjrA1f4JBsWpTKzhEyLYe7ZJSrV9yLMY5kCCppnSVKopbcyg8gqDpGMYyW3Ld4Ot_4nLiWBN8Q4z3AavSvubSi5TyLxbIxMqxYsKnbmH0ls5WjVojp4bm5b0v53lCtsOPXx1js6PT7d9knEpdoPuPeToVAEIhREh7lpbYwLZ6bcXxEQp0KZO2DHJJTpJbKJ5zkKntHn6JyUMuS.cEJdpK6t3GGq0xQQ9M9EyAeB5nYQh4KM6PXsbC54yj2KGbaDzo1MUsTHma0T5a3cXnrToRu4SK2MC1GEZS8Qo.Wbgrj4VzqifhDRIPCtwID4PRqRgtU7mmiTUYdfz8rmGGgiC1kadgKJ_InFRSnNRLr3kBYpt0.KV22_u3OsCIg1UeZW8vlavhP6RWcPOqwrUrbUJPwSYwTuzbY.Ny20Am4DsVVdDyy11166V8YexGxV8eJ7sji29EuHPrsZm3Er9j6TjscjS56jjFMo980K.FjFZI0id0qDOfBGZCI87Qk0MrzzwzTfYMEhIvqoY.sRiMieMUP4HzfXKOiktKffH61NG0AQ0YOxoJYbZGhEchTKYPbKeSX2loSSjSHoRGSNRi8CK_DM8ma8lcHw00YBPAwteC9C.qViDjNlocEh0Iu85bu4K1eboizka_F1rHSWU4UXz1HIdq.JOC9adOzyFQenerw9Zv3GJHaKGy5hgz74fDZOhcpoJcgKJ1w_SVMd3LV9R.r59KtZg84pmvAWx5qMPC_4cBYUJFfSNoyUcZz2AWKtKYqxSChzdhSCyKVb2n6DlHjcT79kGM9m.aMDx04bvsL71GEAzpvhpap7HfBJqtF8l8hcueaHgcYdrm79zE9duKEKhD_hqGAWcj8V31gfpw1U8V0hROiNYmOtZDhqEypU8JVrxQjtgbjjnY6hZKo.oGucVTo1OtCFyxuW3nBZ3NKcxxVgV5P0eBpvdqqQtelv4mbCmmVXMu.uP4Du5fxQYPWism99wq.lV5dsr1CG5DMzk0yqNCYNe_2UQAjn6Q_UYDgMPIKRCWyzYemFHr_Wgx77VCjMWzUvQYVHDFFvYE6BWUt7bMA4_eiLWzhpqLcTjGZAEAdhU7FAupMiV1U1k2jl25iTc2R77QGImjL.8L657I9ddV4v.AB9X6eI460037PFh_s9SU0vzwtYETF.FDqcET0sBgVjzNSDbiGBizozZBIPpP0nJKsPC44Ez6ggHTjDrVtp5xJYBWATi4eVALthZ1XTwy2Z7WbCdgpoCJ0NWe0pUclswY7yd8ZRfWXGV6EQFvQA3GbdyBp6RVDrIKugVmYCodT7qBZysWjTvuNMClvD_iQliR.VzxqFlp6Kz4Enw2ZSSB7knZpn_A2ZlLsO3xZCDD1ntLim7ZDOVjv.psd4zOSCRS4ABwjeg5wQypnNry.1njZJJXC3dwa0kNSWFgR0NEIPMtNTQe5eQZAxi5YGfue7r9pVHKADcV0BE8RwqDmLDMYAfS1ThyUfX9RW56w3QFYbfCtPeK8OnEvfD6YHODiPLy7PhY1gQQ3Y3PDwbpmJic27aQSsyDv1rlEq3xc1fdLTVNEi_Y_U5SqmDgJ6rXKYgErGpcMJYAxD11Mo4cElBQHs2R2WGOstdcqJBV2cSxtc8ayQbzmKuhhcvjOL8lAJdogKXh6_Us0WYPNyoMAOwk9G8K48PydT0B4vW4bJ1oOIe9zh3G5rEBjAEPc.ggUN9HR38zNFsX9zgMFMkmbhiJV03moeGiRtoyeKrK_0Eg5CFnpMr2GYKEsFF9EHokL76fEyiwytSn9GDrQwfmQhs8JYtfKhR3b.wVt8J_sPsdvqdR09cemWFbNr55q9X2MWM9NN4kY.FCKArNSK7O3c5QOt6RSzBWJTOoGimU2SGAUXYAJq1JjuL5EXvtX9x_8xs_9xrC2onDtljDwfSA"};var cpo = document.createElement('script');cpo.src = '/cdn-cgi/challenge-platform/h/b/orchestrate/chl_page/v1?ray=8e9d37886eee9c5d';window._cf_chl_opt.cOgUHash = location.hash === '' && location.href.indexOf('#') !== -1 ? '#' : location.hash;window._cf_chl_opt.cOgUQuery = location.search === '' && location.href.slice(0, location.href.length - window._cf_chl_opt.cOgUHash.length).indexOf('?') !== -1 ? '?' : location.search;if (window.history && window.history.replaceState) {var ogU = location.pathname + window._cf_chl_opt.cOgUQuery + window._cf_chl_opt.cOgUHash;history.replaceState(null, null, "\/doi\/10.1098\/rsbl.2014.0911?__cf_chl_rt_tk=FcKSCv99bfgPY.Za89vGIy05tqG5guaphRU17pHZfKM-1732826886-1.0.1.1-U4AxeuiRHQn27LFznTKugh85disMZF4l4qTzEJhFTEE" + window._cf_chl_opt.cOgUHash);cpo.onload = function() {history.replaceState(null, null, ogU);}}document.getElementsByTagName('head')[0].appendChild(cpo);}());</script></body></html>