CINXE.COM

Just a moment...

<!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,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);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: '9134a43c0e909c83',cH: 'vy.ptJXUL1s8ASy5iSsksCD.xaijFuepMRMN4ju7l88-1739783397-1.2.1.1-jMxDv7JrsghKlSmr2xG0qP9gsAv3wPjWpy4nhWUeIOLCfFYtF2m4alJd.j5Otilw',cUPMDTk: "\/doi\/pdf\/10.1098\/rsos.182138?__cf_chl_tk=G6MX6Xqjb_HYalPGyF6s.3AbkI5z.Fa2jrIqqjHU17E-1739783397-1.0.1.1-nuLS_kSpy1Xtoh8u.swE1kwbXhR3a2hJQLQ90BpkAmc",cFPWv: 'b',cITimeS: '1739783397',cTTimeMs: '1000',cMTimeMs: '390000',cTplC: 0,cTplV: 5,cTplB: 'cf',cK: "",fa: "\/doi\/pdf\/10.1098\/rsos.182138?__cf_chl_f_tk=G6MX6Xqjb_HYalPGyF6s.3AbkI5z.Fa2jrIqqjHU17E-1739783397-1.0.1.1-nuLS_kSpy1Xtoh8u.swE1kwbXhR3a2hJQLQ90BpkAmc",md: "CgmQzaboY.BEM.jf.F6V3h1PyiZjJaL71CKwOdaLQa4-1739783397-1.2.1.1-uPoM3jBZu_pUHWCnp4A5qreEVPb3uqdh5cGBfDqFQbYdLVeJb6yi4v6GTb3k2ygRZoHzciCziV6qSshZ7twGLhPGS4NXCtBjhyl1xY85tXJh1NI_DGU4nK0hiD0Xuz.so1YTsLgJYPcY3xrnkRXE4MenIRxrz_Zj.A5MuywLSSh5pCUxkS5ep0cWeeNOe_aykNzO57Dk0HpSDU3wEHkvlTnc5cm5kUKuwVGf_ijDWtquKerkJpl5Qu1k7VyWnmBVQ8MXcSRbPHSCPT38ISuw88cpjkLerD._giaD..SQT6M192abnh9dJaqtYjxDZe3gUcWZXOAVq5iIXzqyqUp3_5zbTqIi7OO8KnPo5y12mCbWPDeu60A.6zWrprbOmWROr.nlT0VBLNCZLWl_tgJ0tjl0pJp42AZdkLli6hfgvT9SEWBunBzy2bMryABmbs8OkfHii.roEPBZhifjILHNjifHyAV1omHLnztmKPMaEtWLCeA.jD0tXEXLuLw6g0Wvwi7TV8lu92iOXSH0xQ8FDjWdNOaTSxaePEkCR_2tmMkIj8H_OgJZTq1IujVJMWmwyeNUNlEMLuijhMx8tQSK1QVcpE.xQkDpa4sG8U9OpUsVLdPknn3qZb7xT0d4aAJgpWpxgNilJ3aZVI8JZL1E2Ww1v6a2dMDVb9nX_hYzAywkAYVlTmYDvbm2BUcJDd4.QlvwnJF0JhDsAPIP8Z5Pe3qScjSUyV1b1cx_.UfTEk7t8JL3lPdUZ5YIBVblyQfsY7VYjWIapRPL.LpQ3NHtJunbI1ekuRkpoRc4Dyyx9iHDhX8G435bZ3DD05XMSj2GpDuizS1cWV1imRyRuEoA9pzixY6sanqjqYKNnoM.tvRPrQB2NAwFsHwJUQ1yMsHPVcNE6yNcaHMp3Okn2pVMTUrFs_i4J34btACWf_t16iuLHrETeZ_Hnww8pxgMlCa_woug1lyoBLT4QdrAjEojqt8Di1HPPixqy.hIf7MQ9CT5r09EYF7uLHTBbac3Fi6JmbcexgPEKaF8HbTH3sqp5hjb54AvmdmA9v8TgN6a.c.hUck.VDENDB7uIrZHIdXYbUzv4nAXg.PbdhtgzpiOmZdGmwGgbRCxeYYFVmb_lOxLv4Ft1qhHC0YVOA4dqBjmwJFbm7LDynWkblJmIccnW0yzxPvxXVAC5EwGL0_cN_T7ZZmGzJpHH5i_aUEB04THXkXs9negHuolb5T.hSzgbuG1EpT7BI8USJ4dKO_NE77IqbAyIOOz24B2cS4zHTlXU7i3rHrQ0HZBnl21IFA6PcYGXr0.zUh_7pKB7EEkhglLXCk7XxANHSIFZY4Ev4kaKMvAFu._1TN1Qx0OyGpswxs9ZCf1MxbV97Le12zrLojHv3rSBA6XNNLq3KGIkHaZCjzlZILvJqQgL3lXNzbezzgeD44KD4zbLC3Ug5qFVXhZvIeS_qxI_7eY_miNenib3..oVTnrMg8yqkX7onp.9PQBXofLfBWl7XLpUjkJBahrPfmyiVxwvIcsHua5a645N4sw2QLeCZ1jwss8q3cqyrLwmcliBG5slGLFWkCDyAyug2Rou436TuiGwkQPixBuWAfVnpcMQsn8v3B4ohe.ZCAw2KZntpunf.LilMHAhh_dYQoX8se0dg0df_DiBGkIf3M7OayZayKMoULuZbY0jLa1fFQoIeMN9Y1fvtLCbIIUtDsnZwwKF0eveXfNZp6gdXyKRfbYZo9_K61RBnuIdqyM0M8BkLAipqyp0UdFeeZ2DcRnHn3MPQKYLAEslQBtIBoVtk6OkpUIbqtTVd1YGkznU4ciOEVhfDo0BNn.H4huYUy94NiaKfJsK3zViIVYeS7..JJaK3GK1WxM9GlUB.FEOMfFLzEvYaOw.kbg.pOgYBXcBTBrEcrMjbS7bI_1i8WyGm1NohLDULQhbsrdD2UBEB0OUpegAF9i2YXMSnWCkJ6RJ4aMkSJz0YFeq4jUl1lUeBF5T9eMTKJWzTe0aq_JwUWmc5czE8HQaMVFo5Z9_EKpUcDDdOBASSDr9JnYDo0rFFtUysaD4V2s6iKII9tYwT3wvonB9J0Jl54pWlWiTHd5SM1Ex.U0GT86as6QH9arzXcNDExgqvbe7DPrz9eVaR.tNJeHTZCkzrn.uv8LBD_H3Pmce5toh1pZFbQO66f7g1le0CJ7nVhdu9p.zieI7O87zeIFrTgcNP4amCjdOBxGZyFk_JgOtoGHvRxxxfIyfIe8_Q8oF_g.dji3VvfYBf6cB9T539ugO3UKkSNi39J.2H7f9ShdMjzBY1zaIhYs8B1GW2IlI.H3CXUOnP6iRaw04hra_7y0j9_JtQnQMYqfYErWYkp6tJM3pCROaiWn_7fXOLnuJ1BafQhancXLJj.l9_sNh9648hn7Q6NBiI6o2XSYumQEpmU3Z6wkilYzIrj3vbTl_Kj9iEKzh0G1LYSwBBeixbmALvk2zPQSGPLru_rg46jIGHzkyBsZFxY3SAIj29GSKaJYI0KUZjXze780AlC0XmC_et4AtUl4Eiw_pBLClbN87n6d7eHIhYZGyEWNjHUlP7jBvAC.71D_frIN4Y4xy9qwaEv2Hlk",mdrd: "xe5iAX3ybRL.4Ng5zFKHaLUBc2xl1KekVTJvPFsLm0w-1739783397-1.2.1.1-4eR1vQoujmieDPjlDZvd_BKPS0w2PSkXQTCnGjl1ABSrOT2.M4jLf1NVac3_d5l8DGeC3nRyppcWeLMuKJwiwM6GXZ0qZ6b2CuPm73zhhqIe6Z6FqKyZXMRW9mAY4Jrbyz_c0rAp6UmdthheNcsciFo0eRwiVqw4a5vssbob8YFNjDGAPsFQbqmHT9x5k0N_dvw_cu00nAIQYdbIwv2RSpxL3UaR8F2oILbPqhdPQ6mfSE6elohAhU55a7bfvY.2hlti76wpnU9pLiwEZuAoXyIdFVwwDUPyzwd1RaUoiSVG4ziq9P35RslBl8tccqGfaL_zlLeoXHt_6cr8GMv.jHcWihhXBZq5l8IXd3kSR0Te7I1C9Ee46vp9ZJz3t63Uzrim40OSBKFe99oU_q5BevJfu8HDVoWxSoO_wdyoQhik0owsWb.WX8jHyMh_.lMJmbQEQvf9LeXBuseWvaT7mvy2Gq_o5_bTg8GdwY2GViVc1xGzoNTqWsHrbKpDU_2tMm0j.LDS7rLBhNhdpkrbymu56Pp9IhgKrOGIpC7Inrz8MMPz2mni4rLxsZ9icB5uJXPD41IDNsnLiqCpyUYJjMAecXf_IXEhzkGtmV71cbluKgTzMV7ojWy4kBJw7JExmBV.MQ8VuGElBOfrbbynpgbrUTi31cIhsgZx5l_oQFHXHiZHdZsl5Mq6_repiGietEWqYtuWe0PVtkaKFUGPGuUMTm1Ezlid3SaWP03P3uPVOC4g2diKswOTSdqxmjVuJZmh.tDCV6picq5wGfBgpnbOlmRJJlIYo_01Jr4hjfoyNW0I16uQDZc7NREY8WhdUIzclkSM2cJXjRO6YL35tTCODd1XbzcZjiSFbzMGg.3Yg.GPyriVBnnJskSjbByQVNxUue7Dabz6J4m9JxefbALMZBMvjJ0iI6UsBAq5z380r50SX4Zz1_5nlnxWDXq7UsdWnXYWIe2lMi.AxOLNeg2rq01cyKh2BbN2uMKldQfwF9.xpi2coKITf.72XdbRRMBVgvseJZw1M40ziQyzM7RcKUTP9uaiipWA1B0E4QNwTS2XlY4JTn9De_vJRB_0WGkU1WjgRXVTyVq2LP2Qz89lA3R_ES2s6yhotIrpIO5A5u8WYqE2xcqAVrsMmDM4tZQuC7UU8SUz_Yh_VRh9W4iYBoa0pt5Y6qmaks_JNCMaR3vpzB3X9qxlkiKt8x7Jij3MOufgNthOrdf.PNR0VSakDSjGXTvcAr9DGMewFF_UYF.XjjzoaEYUouF.ibOkvMxy2JuPXqMg_ueEvaZtu1JrHduwHQ4YU7JnW2bKE9VGPjZoJTz6EK0FVkYrLZ_RDuaQZYO98hG72HljRAfC9Bpm4IT1n9Q3jkdeJRfkBw9JFzdTFTGejb_lu_7rntc0eS.x6jiWhLVKd510OYnkiom9YGWtD5PxGYxEfvuLuc1yNrD3C9PWXAG8xGJFpxcfKzXI96.7KMr0v87T7j8FqmyTLBJnDDOvWTmpZ6QNB.mD6aHB2vOV4ceJS1WY5qAAj.ATWIl9P.M5V4PHnH93F1vB6qTy.ak0wUswfEMVGnKp.UalT80X9jmjej28lM9Oh36vk2aP_PUZA8YL6pbrAeEr_s9hv4HVU_MbjcBbKRoxgAOC_0Njts0Pz9ot_AUjA1tqFx2sIE40tg7z.icfNwz.qIgwu054J8wrii7pixLaSYUtxyVIw7P0YlxQCGKX.LK0..hdVaPwJ1a4NEL1n.mRsOzlDG5KQ.ruByiWm9T1SPyx3pRi0MsCLtCbGz7HBksL1zrtICqZZzOHAW1sJ_X_RCJQqcqzeX6.u24EsB7ww8ZnPPj2HWGpfJLiN91nfkhER7_TbmAIJ9xHm15ucT2r.e9EhWWNxwWuwnKaXuI5rzYh4jBU9lsUhMPILtAqdS0c3AAvoMxE1zDrJhSdwZ5KjOidVkQKqYc1MjUDYMm0JmFwSFzTpgB0YgYia_O0Q084EoaQ7iJMjpG5CioPfg"};var cpo = document.createElement('script');cpo.src = '/cdn-cgi/challenge-platform/h/b/orchestrate/chl_page/v1?ray=9134a43c0e909c83';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\/pdf\/10.1098\/rsos.182138?__cf_chl_rt_tk=G6MX6Xqjb_HYalPGyF6s.3AbkI5z.Fa2jrIqqjHU17E-1739783397-1.0.1.1-nuLS_kSpy1Xtoh8u.swE1kwbXhR3a2hJQLQ90BpkAmc" + window._cf_chl_opt.cOgUHash);cpo.onload = function() {history.replaceState(null, null, ogU);}}document.getElementsByTagName('head')[0].appendChild(cpo);}());</script></body></html>