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: 'managed',cRay: '91a03ff75f93f8ce',cH: '.mJENZUuhuXy1rN_7y4W5SCaoqI254wyXZ4qQCYmPrY-1740911752-1.2.1.1-z7h7UXQmND1rEmM4iPzA9ArPczHFbroGKx1QFRw2uFyjx7PJgx9MPBO5my.NAZnH',cUPMDTk: "\/botlinnean\/article-lookup\/doi\/10.1111\/j.1095-8339.2009.00996.x?__cf_chl_tk=xOSKvFQyPTuv9ptt49SBIJDBEFAIj42yTuJEImPhz.A-1740911752-1.0.1.1-t6evEfjxCbkAlw1Af79DxD9hwtusLTStpwpQSoDcQPk",cFPWv: 'b',cITimeS: '1740911752',cTTimeMs: '1000',cMTimeMs: '390000',cTplC: 0,cTplV: 5,cTplB: 'cf',cK: "",fa: "\/botlinnean\/article-lookup\/doi\/10.1111\/j.1095-8339.2009.00996.x?__cf_chl_f_tk=xOSKvFQyPTuv9ptt49SBIJDBEFAIj42yTuJEImPhz.A-1740911752-1.0.1.1-t6evEfjxCbkAlw1Af79DxD9hwtusLTStpwpQSoDcQPk",md: "3C1vg2nq_LdYdeZKaqEcRXzFp_ZSckzIv_F3GMvz42g-1740911752-1.2.1.1-qGWbPdRH_qSxj0Fb7VZovqqhzUmKRCNsFRHoKFx7ar664pIscox6Wez1sHdcGk21oonmiOnzA11Dcdk4Lz1AQnrXUj7Jk0x.bAGQ5OZ1qmhL1mnM9_BZ3F5.TtxPw7rARx4BWcbyQwz8Hbd2RTo4iam5Nzh7JGBZLZEsgHXC0dV4qoV04NNwLNaG04BTw26VLONCchiVjnGhkXf6O.di6hOGbjcmHQRhl37OMkyZz6WtIARBOBle_N5I3KRWZN68gxqXeAqgaxHzllpkmhs5wSAp_AVUuqZrSRLfb0hekx6g7o5z1XDVxwIhc.Q8XtO0jPA22p_jWEoI02_AO81B6CnX4SvXaKA5zUJY_AI3J1UQILSWEZgsqPDVq4L4FJyOgV7N15QQzHc9m9tMqpDYlJ5H9PM4MYC15dfkDpPIngfjqie5XYmhRNOyDvJv9V3asaXN5IBrY0uJ.sKmXoM37cdR3a_PBZYvOCesiMBxWkTO90XI6bTPcvj6yy3X6bt4qPoGa4gJtAS9SXymEVZgVzrWe9O7zqQ0N.D55t2hkxjK6KTJ9LPx01fPfrPkOQmMxRVXdesHAwpdraF9yp0nR7lqFzFcYaibGdyUnnenKWzHoYzdnwykmadrHX6nhOOC84i3.sJXy1dfPq3XuoD8emZAOBo9OKXTVWJIUO7EeLEJKb6zo13a0oL3jzO53UHeCSivLz4C.b5yJ8UlHyDoeJMmHwvjC1DupajAVJzUGGC3Mbxdwo0a7tUzN9YsJkU59WWKtUPj8njwF0RY09H6sONHcVYMyqsLf.28AHG9p.swUF5ChrBfoiEEC6TziRXkRAaCi7GNe5VDBi.0jZR6wCD9SL6tbythyQR6cXThy0HQf7agJTosG9y3f5yEmmEvF2XLW0CRJG5CIj15H_hJQqBd6zNJkYNopJbdXgxONHynTn98jk1eCqUnicqdMi.IRpzJPSlX_9ECStyT3CNAZKzmhqqKZtyiKAH5y8vSpMikPBH_TcfIVBWjBx7LTvasEL3oTaddlzjKntZ20vsZ6.9hYQjaPygw1WCvKUEAbc.rBLE4Fxh0423UQ4QN71J1jrZwFx8L5XXAY5QoUIiIc1dsM7OQVxM.JfktwUMV.zQpWk6XaDS8UH95rbbsVh1LPQlX9sCYIGAyx8.mrAB000gcsm_.5JWvAbFUBnuQnXPPMjiEgpceotcqFc6NUWawhlyT65ePpZTwywDeRGFaGc.y6rmMgbn5sua5U9a8qDAu6rQq8Fp24sUTOwwtVu4tvZLzLNoAS2GXpT_RY5OAIVj6pkDFQ9HD9LX_h.UTxA.NIGPH7VrhFQDSqNe.lhZBucqjSQ_NRJZm58_LwB6wb5k0KjYXgEA73EvNIyFYNVo",mdrd: "hqm6Yadv6kicmHkTbUvb.bJOEbdSzI2nPjRBqwWnwuk-1740911752-1.2.1.1-JHensCn9Sfjl9XsGoC1JQ9xmLFgQfslkmA2Gzb2CfhsclE..uI4_qTqafILL0ORevaAuxcH4VEkxD3YsaE9_uELTsfm2JOBs.tJyfm9CRkT5CjLeBx_xRmYElhCqlFo8iIbSWKziFSDRZBjaH7XRPa9ozflSZYkDtog4.hrjaVakQZJj1I7EH0mJipC74wOyGq9TlC8_B2T6ENHTdONZ1fcsI3tDqQrtYquD14iR27l22kNRS4mZBuFjlBPl2GOHa2af.xFj.EJn8y2GznI_Q32g5M7Br_BJQ5V2rnNr6DgsiQgvt_NJ4cwjtntuY8HddHes0B7ou_cMxn1tL4.dwz_iO4VIYD_yHtkSEBk94yLOVzX1zbrgEDKdU9LGFeB16Nb48iq8Aynoa5.3szNL4mdQnywSWkIByX3zkRyrmN7JP5IuWDii4dcgYX5qPnoeFNw2BMs3hlc4U.MVZvr9hFXz1voV4_2cpjD3KO4zir6PkdlAkOBZY4B30iliNK06PVPvmiMDudeD2RPHPGyGoyEdUsNawp0ZAnbf_epiMGbj46QmCuc9tY_cWoBPzOGhJk19sEH3crdwtHhYhYWnFV5sllREyRUlhRNQ9kB6xfVf8DvjdM3p2FhlpNFDmj.gRjAGeq7ME3Qfr4srK9wJRl6KGTB1X0mfgEyPbTEMs6OiDOqZZ6DYjPLykB.FT9Ag.QSSb7_hgmdV2HvFX5.vcseHFkNiQMAVZrYv32FVrmOUcDKkn3E4i5EvXU28nimYJYfMeBGLtO1B5hWb4KVaO2d38R5DvieIRKWzhqiCy5Qg_0URInPqE4BZXwNfWd.tsGE9nqZZ84RzWMtyeoWQWKTFe7xCUmR8jziClj4N_cREHIPhR.fx1aQbQdHno9WHh43WOo2Z7NBM98Zxu9akdSUOXzYAjg8wdYso857DvAl1VcTND0BgKw8nBTKBaETJeu_XlpOC0D2wkv2xMJsjp4FGx0Ae4FhIfteQAsH_S2K_vVRq_p1gX9k7vPJpf5IkqQMG5yIZdUwQ37EDMdvNfMo9LckSbRbV7_q0ZW7bYkl5c_kwhRT40SrLf3TMyPlxsBuAecPg_.MCTR_QRz3TUvx2QFc1zDW3q.GVd1z0VozIWd9lEbAZne9f4ofS2M9qEbK7T7NGvN2g1VQWjetHzFuIX71v8R4.dOAaL14KVV3Y7iv8_BDMWKQtIXSSzrEBUvBxhKSUvENJbi5Yi3MUYgA6os.cj1lTiIV.oTFVlSeaY1LX059BUVxD5rlmWGhkQ8Z5AYFB4FS_UNvN0NVIxPwNz9UV4f7vvP4iFhPhXCbxiIjrLYfdLnKmiiGCTsLp2IRQdWjPPRVbUgFG6IK7YN88RBG5RQ5bo8MqsuljKp.tTUOfW32bOKm.pRaL1UzvNIBMd_qxEuh4sfBLJ39nzAQ1joZh9wk5YjDZ8co.9ndTmncx7JVnnNfP9h_Q7UeuokBpvc3IreFGCiGuEVtbqU_9JdcFbTUOpSiFB4sOxQtC1YNY0qk5g._0H2KVP6pToVnqjXvj7TyjSV_RJ3.IIIzcTPzY3dtTVtCK8ebHeCV2L7hPfAmIY3UzbUnO5xTCdx9ZAB5KNdAQ4Z8sSZcy1B2jfoIsJ3ELhjUqRyluHdZ..3bw_uxBsCY9yP1rJ6cQDxDwX_I03RBNAnc9JCgv6M7GUFaqml2dxEREmk0ABYK0ph00xGnfva_DCj..Dhh5tXdDpnDOuvFcraxbrpKOP.ci1l_xCCyHaap64TiadFL_I2.xmujvOClq9cLGOn_CMT4dKMkkQk84lszg3GWq3C7QgXMIocd6zL2f.ox_PlEfF7HSMl.cavVHED0FJ5F1oRMd7Pw7O9X.SqJ3hueC3fi3oP7jqG8pZvEwRv5ZM9_bslHTqiH2sRjIkguCp8ickEg3DSmMGjmfB3wEBUSWK0VVKuYo4te5VKNkB4LkB1o"};var cpo = document.createElement('script');cpo.src = '/cdn-cgi/challenge-platform/h/b/orchestrate/chl_page/v1?ray=91a03ff75f93f8ce';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, "\/botlinnean\/article-lookup\/doi\/10.1111\/j.1095-8339.2009.00996.x?__cf_chl_rt_tk=xOSKvFQyPTuv9ptt49SBIJDBEFAIj42yTuJEImPhz.A-1740911752-1.0.1.1-t6evEfjxCbkAlw1Af79DxD9hwtusLTStpwpQSoDcQPk" + window._cf_chl_opt.cOgUHash);cpo.onload = function() {history.replaceState(null, null, ogU);}}document.getElementsByTagName('head')[0].appendChild(cpo);}());</script></body></html>