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,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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: "academic.oup.com",cType: 'interactive',cRay: '92696ac8de2d7ac6',cH: 'Pxk.9ACHi4pLZtn_fckwx1Qh9Vibe_cIDzH7Dosa5yU-1743021144-1.2.1.1-O4LfaMlp8hoACHfas.Y96cxtPkyaQzWEUq7vreLvsMWb4HgVrLNWYESWiGibfu80',cUPMDTk: "\/mbe\/article-lookup\/doi\/10.1093\/molbev\/msi027?__cf_chl_tk=wyAzzMvgNEA3gSqU04GsptX2liX9q37V1jZjU7QrAYE-1743021144-1.0.1.1-wM_kc6VL995ZrCKLYfdVjHrNhOxOFXGiwTwmjLaGQ3g",cFPWv: 'b',cITimeS: '1743021144',cTTimeMs: '1000',cMTimeMs: '390000',cTplC: 0,cTplV: 5,cTplB: 'cf',cK: "",fa: "\/mbe\/article-lookup\/doi\/10.1093\/molbev\/msi027?__cf_chl_f_tk=wyAzzMvgNEA3gSqU04GsptX2liX9q37V1jZjU7QrAYE-1743021144-1.0.1.1-wM_kc6VL995ZrCKLYfdVjHrNhOxOFXGiwTwmjLaGQ3g",md: "mQWoDsvzJkAa__YKMrvrha9ypUqmop4HnKfDIwecsnA-1743021144-1.2.1.1-1Q.uxQGwjIdEZr_laHe1POqGoIze_LPtl7rcx31uT5U8Gu0b4wFANMKbf25NwjAw6PYrM_rZpgXnlJcgB2tqu8MF3ww0pGKQxlwmB5HgGp5J1W6ub8L4BaqjEzkb8mEDJ5V8ZHGo2Ovv93LZdg1s.L4Z.YiH.xKf.sKRGdqj4aa4KW2c6maCbA0JzQY2kpPsEpzoky81D.kbYzp0u16Eu2mfPFN1DtwAg0na3O306kL_hylgJRMExp22QVZiH2mAmoide8gDLJf.MqwoKnWJ7eN.qnROOIXdnmlEq9TYPRiTnpBuvympENHYTt1kFAjrGMcjLnPJpT_nr8QyjalQ05iiWpGOTDsqDBRecZ5zw7ur5nEdQiykUunI.fFgBaijhvfw3GR91.t5MAhMFGzMBPhqGVtX.6jfWJhFIltWkDWyjMl60FDuu.RzD6t2d0rUZWZfuf.5NBmBu8XXmo87ObR5QrX.pKx20bbvD99TrNrqcQNLYzs6UvuZlM0hnP2j4BiA6Rc.Ig.zg5e6UHEBsXHYYwda2n2tFhSfc91PyfF3VJLkNzpoPs0SrfRNd48ZXjMPAxHNKONQsyLAOyG1FyL1nqaF9J71Hg1qxN3IJF9SEUQmAzRNP0N3kGcXJwU4vazpr4uNRaSKTeii9AvbY_7UvEOQQbvsBMLcbkp9dsnbCtFEWhat61wvE7x9C2p945eZDu5yL4wzGHrJ2b2cx3CN2gsDy1cNp9XYjxndi2mhfdZ3ObHbYoSK6wJmzsXlLSIuIjV20XkHuHWTd1fVhiqaC.Bbqi_XTNoCsZSnlJU9i8fsvx9kMthuOnaeptuXzerAOdRJjGeJtUp2rpwKirXW4XA7i1nLm1tCO0NjwSL_n_0FQqs.WJz8mxZgd1WrGfYGEI_.Xt.TfS2U5ciegLo3BnpEMCiJ3dWLDRvLp5RIpMb0NaThIee9eLausoqHEyNeF_pkD9SVVJ00m5sULAzv2ArSzfMMRh71ehGA_LQATIJyVs_vKLS_niFiRgzKYnnKsE1yEyhpZk49AhTwtO_Zb6yM9I0KRDmiZUmGALdd8_bYbjvu2klbRxQh4QA6Z6v0vuuFr_P.I71uQvv0_3H6fI1_Nto0NP28NjbrQ7GI4rNVwolgt8UgfDZKU3kw7F1EbLU0L7_ox5cZ4rtLyuIfUXDfeJNAAC3257fD04AW3JejWezVr5h5UPs96eWOnTzcbpdFU8aEMzYMR0dpshRXaU84QzE88Op2p14EUhZz4.gMePYOEUnT__RfVt9NWf0vdkUM2H0tjE8_v0NL1jZnruljUcOTVQDydT8CHqYmwnhTVz9M.FfqIBEsyOP3",mdrd: "nART8G.32NWO8oFXShT5JSgaUNv1mba.8SLAUe9bPdE-1743021144-1.2.1.1-qNoCYpJuXlHs9N31BUCiza6p5pL421EVSuKOWySagyJunMPAlrYBSSSLJfkjDTiv7McHLoDr_tml_al5rP.vYdItd8KmK7O5dtkDJJ.lY40VAdGlbwT55S_Iq.CMyq6WaTtQxrUsnKlhFhdqgNj38jBLXInQwS2qUdIAnHH3aD9YCH358kV8h9183Hi__0w6Mb6RUZObP2ksStNJcT_ivxDl5iSw4gRXC9LBaFAw_vkWqI_OmzsLp.oNQPfWAxb6RgS0CtxSf5up3DBmroLMQYpWclORh2dc0ij1sLH9apinjjrbOW_rKfHo5R36OslAO9PdqihzT.r1Ry02EtckweVMOYQAi6.SN5IlFvb40iDJKeSTuHzxm.fOlsfjJejUbFnpeFYXHAlxv_aIi26zzqH7enKNwMUIAEH8AkCzWmMd4wH2C5wAdrBLmt5GNVKgPceMIUaWGjouIkihPzfgvaFhpsK.qWcTZPU7sAMmoW5QHPHuhJFw22qvTwslulk8FBog6TvuQt0_GfxsP1auv2kR6UHCqeklpxivYq.rBpjEYMwuP9Y4P592lMfnHoAm6VBJAk.ZA7sykPNTMIBDbSjwl5JQyXbPkLrz2vhgqS0gQMv3_At9pf50T1toJ_4ILwLorvKktMg5eb.ORZwVuUi1rRbH16xA_6i16wTGQzaGr46hGk8HbjEcnFHpYr9RQr1fPSiiUs1KnF9btCtpMSUHcWeUuAhD9jktvX01i3T_lpYov5KxPavhfOGno8MqSDv0kdNurlbuSBDhfqlx0XUTOPA12IWboUrxkyVLpP7rFqcnuaBZyZRFSYrdjoCBVMR6Hyu1.nBvq8CiDuCCCOddFrVY0AJYtxeF36tdlacRd8RgSdbwxdEmqQ8fhOnGiVhctcbnl6550esTNMR9CSfAEL2mbimOXhKI74pBearxHaIpWfO5bDfrFzlR_W7xdcl5rQvfJUxEIkmMPuAMyJ_Ki0qL6QAWqr7USnwvz.RFO7MUR0D8DTPrdn8l8Cod_HeY4Q4OBGv0HCllbGOwpHfipis_dG26svnCiXnIRL658kSnuDbmjRTjerJvCb.xK_dfHu.2owGlNUOWtXkoxQlKP7M3TjhGWxhrJKQv79jq.M_cn7MkYdaKZoAhlTVqKsbvJ6N._nNdHg1ZQZzirO_LpQt7dG4j.TLtY0UV4.KbmkbWWPxhx7C2cNHArl6mhV7.1fZV2Op9cRmT.O71CKhIbJ73jnw.ni9QMriUaWULRl0i5O6FlSdgpo_piWyQl1z.NoLxw3V06AtGP9XBXy9smX.YSx9PuC.l3wg_hge00AnAO70CAiDRCW_cyUorydQEAtKzp3J4yhydbEnpcl9blb5_AwVRwoA3RvIEfUejKPGizGx5gucBbdflQK7E5PKTwv2TbWk6.q2ZaX5EyvRRwsmwX8ekL9f5ZQCIQqkUN40vXcGsHXTT3PnlcchOTIlVojyNxcwxb8DufSaQiXxkeZHKNgrMI1wRH_gbs7L.tIGZBE5TW3Y1aVZ_78K7ddcj_lR9DUaRNvn0_wns1GGKydA_gEV3wHLFIRp44k8LWVJSWloOeIeE4v2RypGZe6EYzWqVihzAjvmnJvFMNZ2g02Yth1iyjP2TILf9sRaoLCVggUr6Pk3o3br0oOiOTS6BWlZbOsXC4hnWOcgOOM7zFoYdv4PA7zmh77n.AZKhjI_3Ir1mgmoW.tc3Omzc4SllbnVH1acvSMwe8SFa7mpL4kqTWJucsVwMBjZQzaSOMb0LVFyJk8grM16NVs1MNGOpU8aHZecPUTz6_KN.wh.oOlPpx6pqTVPfcZ31BgxWMq7FxfNcaawx..FLjvujnyGiy8geKd4NEmWx11koWECbGuFjacGG.a6DL3VydYcVcHG2V.hD_dGUWBeXfRFfj57mlp_1OCeHgBSL9k7zBNGG3.oh1nLp9_dyWE0BontAX270zXWgHy_vimwZBzDLu0GsAlM8FC4Oayl6z29O01SVTuC8Xn8Ur3LPICaZqWdyW_NcUgjP1dh2MyKGjJh4S8xGY6QUjHeK36GAoXMEhKewvC_to9iao_eHZicYWWA"};var cpo = document.createElement('script');cpo.src = '/cdn-cgi/challenge-platform/h/b/orchestrate/chl_page/v1?ray=92696ac8de2d7ac6';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, "\/mbe\/article-lookup\/doi\/10.1093\/molbev\/msi027?__cf_chl_rt_tk=wyAzzMvgNEA3gSqU04GsptX2liX9q37V1jZjU7QrAYE-1743021144-1.0.1.1-wM_kc6VL995ZrCKLYfdVjHrNhOxOFXGiwTwmjLaGQ3g" + window._cf_chl_opt.cOgUHash);cpo.onload = function() {history.replaceState(null, null, ogU);}}document.getElementsByTagName('head')[0].appendChild(cpo);}());</script></body></html>