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<!DOCTYPE html> <html lang="en" class="grade-c"> <head> <title>The hornwort genome and early land plant evolution | Nature Plants</title> <link rel="alternate" type="application/rss+xml" href="https://www.nature.com/nplants.rss"/> <script id="save-data-connection-testing"> function hasConnection() { return navigator.connection || navigator.mozConnection || navigator.webkitConnection || navigator.msConnection; } function createLink(src) { var preloadLink = document.createElement("link"); preloadLink.rel = "preload"; preloadLink.href = src; preloadLink.as = "font"; preloadLink.type = "font/woff2"; preloadLink.crossOrigin = ""; document.head.insertBefore(preloadLink, document.head.firstChild); } var connectionDetail = { saveDataEnabled: false, slowConnection: false }; var connection = hasConnection(); if (connection) { connectionDetail.saveDataEnabled = connection.saveData; if (/\slow-2g|2g/.test(connection.effectiveType)) { connectionDetail.slowConnection = true; } } if 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Here, we report the draft genome sequence of the hornwort Anthoceros angustus. Phylogenomic inferences confirm the monophyly of bryophytes, with hornworts sister to liverworts and mosses. The simple morphology of hornworts correlates with low genetic redundancy in plant body plan, while the basic transcriptional regulation toolkit for plant development has already been established in this early land plant lineage. Although the Anthoceros genome is small and characterized by minimal redundancy, expansions are observed in gene families related to RNA editing, UV protection and desiccation tolerance. The genome of A. angustus bears the signatures of horizontally transferred genes from bacteria and fungi, in particular of genes operating in stress-response and metabolic pathways. Our study provides insight into the unique features of hornworts and their molecular adaptations to live on land. A draft genome sequence of the hornwort Anthoceros augustus confirms the phylogenetic relationships among the three clades of bryophytes and provides insight into the unique characteristics of hornworts and their adaptations to live on land.","datePublished":"2020-02-10T00:00:00Z","dateModified":"2020-02-10T00:00:00Z","pageStart":"107","pageEnd":"118","license":"http://creativecommons.org/licenses/by/4.0/","sameAs":"https://doi.org/10.1038/s41477-019-0588-4","keywords":["Comparative genomics","Phylogenetics","Plant evolution","Life Sciences","general","Plant 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class="c-article-author-list c-article-author-list--long" data-test="authors-list" data-component-authors-activator="authors-list"><li class="c-article-author-list__item"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Jian-Zhang-Aff1" data-author-popup="auth-Jian-Zhang-Aff1" data-author-search="Zhang, Jian">Jian Zhang</a><sup class="u-js-hide"><a href="#Aff1">1</a></sup><sup class="u-js-hide"> <a href="#na1">na1</a></sup>, </li><li class="c-article-author-list__item"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Xin_Xing-Fu-Aff1-Aff2" data-author-popup="auth-Xin_Xing-Fu-Aff1-Aff2" data-author-search="Fu, Xin-Xing">Xin-Xing Fu</a><sup class="u-js-hide"><a href="#Aff1">1</a>,<a href="#Aff2">2</a></sup><sup class="u-js-hide"> <a href="#na1">na1</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Rui_Qi-Li-Aff1" data-author-popup="auth-Rui_Qi-Li-Aff1" data-author-search="Li, Rui-Qi">Rui-Qi Li</a><sup class="u-js-hide"><a href="#Aff1">1</a></sup><sup class="u-js-hide"> <a href="#na1">na1</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Xiang-Zhao-Aff3" data-author-popup="auth-Xiang-Zhao-Aff3" data-author-search="Zhao, Xiang">Xiang Zhao</a><sup class="u-js-hide"><a href="#Aff3">3</a></sup><sup class="u-js-hide"> <a href="#na1">na1</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Yang-Liu-Aff4-Aff5" data-author-popup="auth-Yang-Liu-Aff4-Aff5" data-author-search="Liu, Yang">Yang Liu</a><sup class="u-js-hide"><a href="#Aff4">4</a>,<a href="#Aff5">5</a></sup><sup class="u-js-hide"> <a href="#na1">na1</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Ming_He-Li-Aff6" data-author-popup="auth-Ming_He-Li-Aff6" data-author-search="Li, Ming-He">Ming-He Li</a><sup class="u-js-hide"><a href="#Aff6">6</a></sup><sup class="u-js-hide"> <a href="#na1">na1</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Arthur-Zwaenepoel-Aff7-Aff8" data-author-popup="auth-Arthur-Zwaenepoel-Aff7-Aff8" data-author-search="Zwaenepoel, Arthur">Arthur Zwaenepoel</a><sup class="u-js-hide"><a href="#Aff7">7</a>,<a href="#Aff8">8</a></sup><sup class="u-js-hide"> <a href="#na1">na1</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Hong-Ma-Aff9" data-author-popup="auth-Hong-Ma-Aff9" data-author-search="Ma, Hong">Hong Ma</a><span class="u-js-hide"> <a class="js-orcid" href="http://orcid.org/0000-0001-8717-4422"><span class="u-visually-hidden">ORCID: </span>orcid.org/0000-0001-8717-4422</a></span><sup class="u-js-hide"><a href="#Aff9">9</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Bernard-Goffinet-Aff10" data-author-popup="auth-Bernard-Goffinet-Aff10" data-author-search="Goffinet, Bernard">Bernard Goffinet</a><span class="u-js-hide"> <a class="js-orcid" href="http://orcid.org/0000-0002-2754-3895"><span class="u-visually-hidden">ORCID: </span>orcid.org/0000-0002-2754-3895</a></span><sup class="u-js-hide"><a href="#Aff10">10</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Yan_Long-Guan-Aff11" data-author-popup="auth-Yan_Long-Guan-Aff11" data-author-search="Guan, Yan-Long">Yan-Long Guan</a><sup class="u-js-hide"><a href="#Aff11">11</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Jia_Yu-Xue-Aff12" data-author-popup="auth-Jia_Yu-Xue-Aff12" data-author-search="Xue, Jia-Yu">Jia-Yu Xue</a><sup class="u-js-hide"><a href="#Aff12">12</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Yi_Ying-Liao-Aff4-Aff13" data-author-popup="auth-Yi_Ying-Liao-Aff4-Aff13" data-author-search="Liao, Yi-Ying">Yi-Ying Liao</a><sup class="u-js-hide"><a href="#Aff4">4</a>,<a href="#Aff13">13</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Qing_Feng-Wang-Aff13" data-author-popup="auth-Qing_Feng-Wang-Aff13" data-author-search="Wang, Qing-Feng">Qing-Feng Wang</a><span class="u-js-hide"> <a class="js-orcid" href="http://orcid.org/0000-0001-9143-8849"><span class="u-visually-hidden">ORCID: </span>orcid.org/0000-0001-9143-8849</a></span><sup class="u-js-hide"><a href="#Aff13">13</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Qing_Hua-Wang-Aff1" data-author-popup="auth-Qing_Hua-Wang-Aff1" data-author-search="Wang, Qing-Hua">Qing-Hua Wang</a><sup class="u-js-hide"><a href="#Aff1">1</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Jie_Yu-Wang-Aff6-Aff14" data-author-popup="auth-Jie_Yu-Wang-Aff6-Aff14" data-author-search="Wang, Jie-Yu">Jie-Yu Wang</a><sup class="u-js-hide"><a href="#Aff6">6</a>,<a href="#Aff14">14</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Guo_Qiang-Zhang-Aff6" data-author-popup="auth-Guo_Qiang-Zhang-Aff6" data-author-search="Zhang, Guo-Qiang">Guo-Qiang Zhang</a><span class="u-js-hide"> <a class="js-orcid" href="http://orcid.org/0000-0001-6439-019X"><span class="u-visually-hidden">ORCID: </span>orcid.org/0000-0001-6439-019X</a></span><sup class="u-js-hide"><a href="#Aff6">6</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Zhi_Wen-Wang-Aff3" data-author-popup="auth-Zhi_Wen-Wang-Aff3" data-author-search="Wang, Zhi-Wen">Zhi-Wen Wang</a><sup class="u-js-hide"><a href="#Aff3">3</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Yu-Jia-Aff1" data-author-popup="auth-Yu-Jia-Aff1" data-author-search="Jia, Yu">Yu Jia</a><sup class="u-js-hide"><a href="#Aff1">1</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Mei_Zhi-Wang-Aff1" data-author-popup="auth-Mei_Zhi-Wang-Aff1" data-author-search="Wang, Mei-Zhi">Mei-Zhi Wang</a><sup class="u-js-hide"><a href="#Aff1">1</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Shan_Shan-Dong-Aff4" data-author-popup="auth-Shan_Shan-Dong-Aff4" data-author-search="Dong, Shan-Shan">Shan-Shan Dong</a><sup class="u-js-hide"><a href="#Aff4">4</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Jian_Fen-Yang-Aff4" data-author-popup="auth-Jian_Fen-Yang-Aff4" data-author-search="Yang, Jian-Fen">Jian-Fen Yang</a><sup class="u-js-hide"><a href="#Aff4">4</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Yuan_Nian-Jiao-Aff1" data-author-popup="auth-Yuan_Nian-Jiao-Aff1" data-author-search="Jiao, Yuan-Nian">Yuan-Nian Jiao</a><span class="u-js-hide"> <a class="js-orcid" href="http://orcid.org/0000-0002-8987-2782"><span class="u-visually-hidden">ORCID: </span>orcid.org/0000-0002-8987-2782</a></span><sup class="u-js-hide"><a href="#Aff1">1</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Ya_Long-Guo-Aff1" data-author-popup="auth-Ya_Long-Guo-Aff1" data-author-search="Guo, Ya-Long">Ya-Long Guo</a><span class="u-js-hide"> <a class="js-orcid" href="http://orcid.org/0000-0002-4643-4889"><span class="u-visually-hidden">ORCID: </span>orcid.org/0000-0002-4643-4889</a></span><sup class="u-js-hide"><a href="#Aff1">1</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Hong_Zhi-Kong-Aff1" data-author-popup="auth-Hong_Zhi-Kong-Aff1" data-author-search="Kong, Hong-Zhi">Hong-Zhi Kong</a><span class="u-js-hide"> <a class="js-orcid" href="http://orcid.org/0000-0002-0034-0510"><span class="u-visually-hidden">ORCID: </span>orcid.org/0000-0002-0034-0510</a></span><sup class="u-js-hide"><a href="#Aff1">1</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-An_Ming-Lu-Aff1" data-author-popup="auth-An_Ming-Lu-Aff1" data-author-search="Lu, An-Ming">An-Ming Lu</a><sup class="u-js-hide"><a href="#Aff1">1</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Huan_Ming-Yang-Aff5" data-author-popup="auth-Huan_Ming-Yang-Aff5" data-author-search="Yang, Huan-Ming">Huan-Ming Yang</a><sup class="u-js-hide"><a href="#Aff5">5</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Shou_Zhou-Zhang-Aff4" data-author-popup="auth-Shou_Zhou-Zhang-Aff4" data-author-search="Zhang, Shou-Zhou" data-corresp-id="c1">Shou-Zhou Zhang<svg width="16" height="16" focusable="false" role="img" aria-hidden="true" class="u-icon"><use xmlns:xlink="http://www.w3.org/1999/xlink" xlink:href="#icon-eds-i-mail-medium"></use></svg></a><span class="u-js-hide"> <a class="js-orcid" href="http://orcid.org/0000-0001-9070-0593"><span class="u-visually-hidden">ORCID: </span>orcid.org/0000-0001-9070-0593</a></span><sup class="u-js-hide"><a href="#Aff4">4</a></sup><sup class="u-js-hide"> <a href="#na2">na2</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Yves-Van_de_Peer-Aff7-Aff8-Aff15-Aff16" data-author-popup="auth-Yves-Van_de_Peer-Aff7-Aff8-Aff15-Aff16" data-author-search="Van de Peer, Yves" data-corresp-id="c2">Yves Van de Peer<svg width="16" height="16" focusable="false" role="img" aria-hidden="true" class="u-icon"><use xmlns:xlink="http://www.w3.org/1999/xlink" xlink:href="#icon-eds-i-mail-medium"></use></svg></a><span class="u-js-hide"> <a class="js-orcid" href="http://orcid.org/0000-0003-4327-3730"><span class="u-visually-hidden">ORCID: </span>orcid.org/0000-0003-4327-3730</a></span><sup class="u-js-hide"><a href="#Aff7">7</a>,<a href="#Aff8">8</a>,<a href="#Aff15">15</a>,<a href="#Aff16">16</a></sup><sup class="u-js-hide"> <a href="#na2">na2</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Zhong_Jian-Liu-Aff6-Aff14-Aff17" data-author-popup="auth-Zhong_Jian-Liu-Aff6-Aff14-Aff17" data-author-search="Liu, Zhong-Jian" data-corresp-id="c3">Zhong-Jian Liu<svg width="16" height="16" focusable="false" role="img" aria-hidden="true" class="u-icon"><use xmlns:xlink="http://www.w3.org/1999/xlink" xlink:href="#icon-eds-i-mail-medium"></use></svg></a><span class="u-js-hide"> <a class="js-orcid" href="http://orcid.org/0000-0003-4390-3878"><span class="u-visually-hidden">ORCID: </span>orcid.org/0000-0003-4390-3878</a></span><sup class="u-js-hide"><a href="#Aff6">6</a>,<a href="#Aff14">14</a>,<a href="#Aff17">17</a></sup><sup class="u-js-hide"> <a href="#na2">na2</a></sup> & </li><li class="c-article-author-list__show-more" aria-label="Show all 30 authors for this article" title="Show all 30 authors for this article">…</li><li class="c-article-author-list__item"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Zhi_Duan-Chen-Aff1-Aff13" data-author-popup="auth-Zhi_Duan-Chen-Aff1-Aff13" data-author-search="Chen, Zhi-Duan" data-corresp-id="c4">Zhi-Duan Chen<svg width="16" height="16" focusable="false" role="img" aria-hidden="true" class="u-icon"><use xmlns:xlink="http://www.w3.org/1999/xlink" xlink:href="#icon-eds-i-mail-medium"></use></svg></a><span class="u-js-hide"> <a class="js-orcid" href="http://orcid.org/0000-0002-7716-4834"><span class="u-visually-hidden">ORCID: </span>orcid.org/0000-0002-7716-4834</a></span><sup class="u-js-hide"><a href="#Aff1">1</a>,<a href="#Aff13">13</a></sup><sup class="u-js-hide"> <a href="#na2">na2</a></sup> </li></ul><button aria-expanded="false" class="c-article-author-list__button"><svg width="16" height="16" focusable="false" role="img" aria-hidden="true" class="u-icon"><use xmlns:xlink="http://www.w3.org/1999/xlink" xlink:href="#icon-eds-i-chevron-down-medium"></use></svg><span>Show authors</span></button> <p class="c-article-info-details" data-container-section="info"> <a data-test="journal-link" href="/nplants" data-track="click" data-track-action="journal homepage" data-track-category="article body" data-track-label="link"><i data-test="journal-title">Nature Plants</i></a> <b data-test="journal-volume"><span class="u-visually-hidden">volume</span> 6</b>, <span class="u-visually-hidden">pages </span>107–118 (<span data-test="article-publication-year">2020</span>)<a href="#citeas" class="c-article-info-details__cite-as u-hide-print" data-track="click" data-track-action="cite this article" data-track-label="link">Cite this article</a> </p> <div class="c-article-metrics-bar__wrapper u-clear-both"> <ul class="c-article-metrics-bar u-list-reset"> <li class=" c-article-metrics-bar__item" data-test="access-count"> <p class="c-article-metrics-bar__count">39k <span class="c-article-metrics-bar__label">Accesses</span></p> </li> <li class="c-article-metrics-bar__item" data-test="altmetric-score"> <p class="c-article-metrics-bar__count">90 <span class="c-article-metrics-bar__label">Altmetric</span></p> </li> <li class="c-article-metrics-bar__item"> <p class="c-article-metrics-bar__details"><a href="/articles/s41477-019-0588-4/metrics" data-track="click" data-track-action="view metrics" data-track-label="link" rel="nofollow">Metrics <span class="u-visually-hidden">details</span></a></p> </li> </ul> </div> </header> <div class="u-js-hide" data-component="article-subject-links"> <h3 class="c-article__sub-heading">Subjects</h3> <ul class="c-article-subject-list"> <li class="c-article-subject-list__subject"><a href="/subjects/comparative-genomics" data-track="click" data-track-action="view subject" data-track-label="link">Comparative genomics</a></li><li class="c-article-subject-list__subject"><a href="/subjects/phylogenetics" data-track="click" data-track-action="view subject" data-track-label="link">Phylogenetics</a></li><li class="c-article-subject-list__subject"><a href="/subjects/plant-evolution" data-track="click" data-track-action="view subject" data-track-label="link">Plant evolution</a></li> </ul> </div> </div> <div class="c-article-body"> <section aria-labelledby="Abs1" data-title="Abstract" lang="en"><div class="c-article-section" id="Abs1-section"><h2 class="c-article-section__title js-section-title js-c-reading-companion-sections-item" id="Abs1">Abstract</h2><div class="c-article-section__content" id="Abs1-content"><p>Hornworts, liverworts and mosses are three early diverging clades of land plants, and together comprise the bryophytes. Here, we report the draft genome sequence of the hornwort <i>Anthoceros angustus</i>. Phylogenomic inferences confirm the monophyly of bryophytes, with hornworts sister to liverworts and mosses. The simple morphology of hornworts correlates with low genetic redundancy in plant body plan, while the basic transcriptional regulation toolkit for plant development has already been established in this early land plant lineage. Although the <i>Anthoceros</i> genome is small and characterized by minimal redundancy, expansions are observed in gene families related to RNA editing, UV protection and desiccation tolerance. The genome of <i>A. angustus</i> bears the signatures of horizontally transferred genes from bacteria and fungi, in particular of genes operating in stress-response and metabolic pathways. Our study provides insight into the unique features of hornworts and their molecular adaptations to live on land.</p></div></div></section> <section aria-labelledby="inline-recommendations" data-title="Inline Recommendations" class="c-article-recommendations" data-track-component="inline-recommendations"> <h3 class="c-article-recommendations-title" id="inline-recommendations">Similar content being viewed by others</h3> <div class="c-article-recommendations-list"> <div class="c-article-recommendations-list__item"> <article class="c-article-recommendations-card" itemscope itemtype="http://schema.org/ScholarlyArticle"> <div class="c-article-recommendations-card__img"><img src="https://media.springernature.com/w215h120/springer-static/image/art%3A10.1038%2Fs41477-020-0618-2/MediaObjects/41477_2020_618_Fig1_HTML.png" loading="lazy" alt=""></div> <div class="c-article-recommendations-card__main"> <h3 class="c-article-recommendations-card__heading" itemprop="name headline"> <a class="c-article-recommendations-card__link" itemprop="url" href="https://www.nature.com/articles/s41477-020-0618-2?fromPaywallRec=false" data-track="select_recommendations_1" data-track-context="inline recommendations" data-track-action="click recommendations inline - 1" data-track-label="10.1038/s41477-020-0618-2"><i>Anthoceros</i> genomes illuminate the origin of land plants and the unique biology of hornworts </a> </h3> <div class="c-article-meta-recommendations" data-test="recommendation-info"> <span class="c-article-meta-recommendations__item-type">Article</span> <span class="c-article-meta-recommendations__access-type">Open access</span> <span class="c-article-meta-recommendations__date">13 March 2020</span> </div> </div> </article> </div> <div class="c-article-recommendations-list__item"> <article class="c-article-recommendations-card" itemscope itemtype="http://schema.org/ScholarlyArticle"> <div class="c-article-recommendations-card__img"><img src="https://media.springernature.com/w215h120/springer-static/image/art%3A10.1038%2Fs41477-024-01883-w/MediaObjects/41477_2024_1883_Fig1_HTML.png" loading="lazy" alt=""></div> <div class="c-article-recommendations-card__main"> <h3 class="c-article-recommendations-card__heading" itemprop="name headline"> <a class="c-article-recommendations-card__link" itemprop="url" href="https://www.nature.com/articles/s41477-024-01883-w?fromPaywallRec=false" data-track="select_recommendations_2" data-track-context="inline recommendations" data-track-action="click recommendations inline - 2" data-track-label="10.1038/s41477-024-01883-w">Pan-phylum genomes of hornworts reveal conserved autosomes but dynamic accessory and sex chromosomes </a> </h3> <div class="c-article-meta-recommendations" data-test="recommendation-info"> <span class="c-article-meta-recommendations__item-type">Article</span> <span class="c-article-meta-recommendations__date">03 January 2025</span> </div> </div> </article> </div> <div class="c-article-recommendations-list__item"> <article class="c-article-recommendations-card" itemscope itemtype="http://schema.org/ScholarlyArticle"> <div class="c-article-recommendations-card__img"><img src="https://media.springernature.com/w215h120/springer-static/image/art%3A10.1038%2Fs41467-022-31280-w/MediaObjects/41467_2022_31280_Fig1_HTML.png" loading="lazy" alt=""></div> <div class="c-article-recommendations-card__main"> <h3 class="c-article-recommendations-card__heading" itemprop="name headline"> <a class="c-article-recommendations-card__link" itemprop="url" href="https://www.nature.com/articles/s41467-022-31280-w?fromPaywallRec=false" data-track="select_recommendations_3" data-track-context="inline recommendations" data-track-action="click recommendations inline - 3" data-track-label="10.1038/s41467-022-31280-w">The genomic basis of the plant island syndrome in Darwin’s giant daisies </a> </h3> <div class="c-article-meta-recommendations" data-test="recommendation-info"> <span class="c-article-meta-recommendations__item-type">Article</span> <span class="c-article-meta-recommendations__access-type">Open access</span> <span class="c-article-meta-recommendations__date">28 June 2022</span> </div> </div> </article> </div> </div> </section> <script> window.dataLayer = window.dataLayer || []; window.dataLayer.push({ recommendations: { recommender: 'semantic', model: 'specter', policy_id: 'NA', timestamp: 1739724191, embedded_user: 'null' } }); </script> <div class="main-content"> <section data-title="Main"><div class="c-article-section" id="Sec1-section"><h2 class="c-article-section__title js-section-title js-c-reading-companion-sections-item" id="Sec1">Main</h2><div class="c-article-section__content" id="Sec1-content"><p>Land plants (Embryophyta) probably originated in the early Palaeozoic<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 1" title="Puttick, M. N. et al. The interrelationships of land plants and the nature of the ancestral embryophyte. Curr. Biol. 28, 733–745 (2018)." href="/articles/s41477-019-0588-4#ref-CR1" id="ref-link-section-d326961673e997">1</a></sup>, initiating the colonization of the terrestrial habitat. Because bryophytes (hornworts, liverworts and mosses) emerged from the early split in the diversification of land plants, they are key to the study of early land plant evolution (Supplementary Note <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">1.1</a>). Unlike other extant land plants, the vegetative body of bryophytes is the haploid gametophyte, the sporophyte is always unbranched and permanently attached to the maternal plant, and both generations lack lignified vascular tissue<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2" title="Goffinet, B. & Buck, W. R. in The Evolution of Plant Form (eds Ambrose, B. & Purruganan, M.) 51–90 (Wiley–Blackwell, 2013)." href="/articles/s41477-019-0588-4#ref-CR2" id="ref-link-section-d326961673e1004">2</a></sup>. Bryophytes occur in nearly all terrestrial habitats on all continents but are absent from marine environments<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 3" title="von Konrat, M., Shaw, A. J. & Renzaglia, K. S. A special issue of Phytotaxa dedicated to bryophytes: the closest living relatives of early land plants. Phytotaxa 9, 5–10 (2010)." href="/articles/s41477-019-0588-4#ref-CR3" id="ref-link-section-d326961673e1008">3</a></sup>.</p><p>With only 200–250 species worldwide, the diversity of hornworts is much lower than that of the other six extant lineages of embryophytes (angiosperms, gymnosperms, ferns, lycophytes, mosses and liverworts)<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 4" title="Christenhusz, M. J. M. & Byng, J. W. The number of known plants species in the world and its annual increase. Phytotaxa 261, 201–217 (2016)." href="/articles/s41477-019-0588-4#ref-CR4" id="ref-link-section-d326961673e1015">4</a></sup>. Long considered sister to all other land plants, or sister to all extant vascular plants, hornworts have recently been resolved as sister to the setaphytes (that is, the mosses and liverworts) within monophyletic bryophytes<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 1" title="Puttick, M. N. et al. The interrelationships of land plants and the nature of the ancestral embryophyte. Curr. Biol. 28, 733–745 (2018)." href="/articles/s41477-019-0588-4#ref-CR1" id="ref-link-section-d326961673e1019">1</a>,<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" title="Qiu, Y. L. et al. The deepest divergences in land plants inferred from phylogenomic evidence. Proc. Natl Acad. Sci. USA 103, 15511–15516 (2006)." href="#ref-CR5" id="ref-link-section-d326961673e1022">5</a>,<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" title="Wickett, N. J. et al. Phylotranscriptomic analysis of the origin and early diversification of land plants. Proc. Natl Acad. Sci. USA 111, E4859–E4868 (2014)." href="#ref-CR6" id="ref-link-section-d326961673e1022_1">6</a>,<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" title="Cox, C. J., Li, B., Foster, P. G., Embley, T. M. & Civan, P. Conflicting phylogenies for early land plants are caused by composition biases among synonymous substitutions. Syst. Biol. 63, 272–279 (2014)." href="#ref-CR7" id="ref-link-section-d326961673e1022_2">7</a>,<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 8" title="Liu, Y., Cox, C. J., Wang, W. & Goffinet, B. Mitochondrial phylogenomics of early land plants: mitigating the effects of saturation, compositional heterogeneity, and codon-usage bias. Syst. Biol. 63, 862–878 (2014)." href="/articles/s41477-019-0588-4#ref-CR8" id="ref-link-section-d326961673e1025">8</a></sup>. Still, hornworts possess a series of distinct features<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 9" title="Villarreal, J. C. & Renzaglia, K. S. The hornworts: important advancements in early land plant evolution. J. Bryol. 37, 157–170 (2015)." href="/articles/s41477-019-0588-4#ref-CR9" id="ref-link-section-d326961673e1029">9</a></sup>. For instance, most hornworts have chloroplasts with CO<sub>2</sub>-concentrating pyrenoids, which have not been found in any other land plants but are widespread among green algae<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 10" title="Villarreal, J. C. & Renner, S. S. Hornwort pyrenoids, carbon-concentrating structures, evolved and were lost at least five times during the last 100 million years. Proc. Natl Acad. Sci. USA 109, 18873–18878 (2012)." href="/articles/s41477-019-0588-4#ref-CR10" id="ref-link-section-d326961673e1035">10</a></sup>. Other unusual features of hornworts include the persistent basal meristem in the sporophyte and mucilage-filled cavities for colonial symbionts on the gametophyte<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 11" title="Renzaglia, K. S., Villarreal, J. C. & Duff, R. J. in Bryophyte Biology (eds Goffinet, B. & Shaw, J.) 139–171 (Cambridge Univ. Press, 2009)." href="/articles/s41477-019-0588-4#ref-CR11" id="ref-link-section-d326961673e1040">11</a></sup>. Most hornworts form tight symbiotic relationships with cyanobacteria<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 12" title="Adams, D. G. & Duggan, P. S. Cyanobacteria–bryophyte symbioses. J. Exp. Bot. 59, 1047–1058 (2008)." href="/articles/s41477-019-0588-4#ref-CR12" id="ref-link-section-d326961673e1044">12</a></sup> and fungal endophytes (especially Glomeromycota and Mucoromycotina)<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 13" title="Desirὸ, A., Duckett, J. G., Pressel, S., Villarreal, J. C. & Bidartondo, M. I. Fungal symbioses in hornworts: a chequered history. Proc. R. Soc. B 280, 1759 (2013)." href="/articles/s41477-019-0588-4#ref-CR13" id="ref-link-section-d326961673e1048">13</a></sup>.</p><p>Here, we present the draft genome of <i>A. angustus</i> Steph. (Anthocerotaceae) (see Methods, Supplementary Figs. <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">1</a> and <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">2</a>, and Supplementary Note <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">1.2</a>). Completion of this high-quality hornwort genome complements previously sequenced representatives of the mosses (<i>Physcomitrella patens</i><sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 14" title="Rensing, S. A. et al. The Physcomitrella genome reveals evolutionary insights into the conquest of land by plants. Science 319, 64–69 (2008)." href="/articles/s41477-019-0588-4#ref-CR14" id="ref-link-section-d326961673e1069">14</a></sup>) and liverworts (<i>Marchantia polymorpha</i><sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 15" title="Bowman, J. L. et al. Insights into land plant evolution garnered from the Marchantia polymorpha genome. Cell 171, 287–304 (2017)." href="/articles/s41477-019-0588-4#ref-CR15" id="ref-link-section-d326961673e1076">15</a></sup>) and provides a unique opportunity to revisit bryophyte phylogeny, early land plant evolution and the adaptation of plants to live on land.</p></div></div></section><section data-title="Genome assembly and annotation"><div class="c-article-section" id="Sec2-section"><h2 class="c-article-section__title js-section-title js-c-reading-companion-sections-item" id="Sec2">Genome assembly and annotation</h2><div class="c-article-section__content" id="Sec2-content"><p>We sequenced the genome of <i>A. angustus</i> (a single individual of unknown sex from the dioecious species) using a combination of Illumina and Oxford Nanopore high-throughput sequencing systems (see Methods). We generated 126.53 Gb raw reads from Illumina and 63.61 Gb raw reads from Nanopore sequencing platforms, and retained 17.10 Gb and 3.78 Gb, respectively, after filtering, error-correction and decontamination (see Methods, Supplementary Figs. <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">2</a>–<a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">4</a> and Supplementary Tables <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">1</a>–<a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">3</a>). Finally, we obtained an optimized assembly of 119 Mb with a contig N50 length of 796.64 kb and a scaffold N50 length of 1.09 Mb (Table <a data-track="click" data-track-label="link" data-track-action="table anchor" href="/articles/s41477-019-0588-4#Tab1">1</a> and Supplementary Table <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">4</a>). Approximately 97.66% of the vegetative gametophyte transcriptome data for <i>A. angustus</i> genome annotation can be mapped to the assembled genome (Supplementary Table <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">5</a>). Repeat sequences comprise 64.21% of the assembled genome, with transposable elements (TEs) being the major component (Table <a data-track="click" data-track-label="link" data-track-action="table anchor" href="/articles/s41477-019-0588-4#Tab1">1</a> and Supplementary Tables <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">6</a> and <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">7</a>). Among the TEs, long terminal repeats (LTRs) are the most abundant (Supplementary Table <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">7</a>). We used a combination of de novo, homology-based and RNA sequence-based predictions to obtain gene models for the <i>A. angustus</i> genome (Supplementary Table <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">8</a>). In total, we predicted 14,629 protein-coding genes with an average coding-sequence length of 1.31 kb and an average of 4.81 exons per gene (Table <a data-track="click" data-track-label="link" data-track-action="table anchor" href="/articles/s41477-019-0588-4#Tab1">1</a>, Supplementary Fig. <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">5</a> and Supplementary Table <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">8</a>). About 85% of these predicted genes have their best hits on plant sequences from the National Center for Biotechnology Information (NCBI) non-redundant database (Supplementary Fig. <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">6</a>), and 78.39% were functionally annotated through Swissprot, TrEMBL, Pfam, gene ontology (GO) and Kyoto Encyclopedia of Genes and Genomes (KEGG) (Supplementary Tables <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">9</a> and <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM3">32</a>). Our annotation captured 89.64% of the 956 genes in the BUSCO plantae dataset<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 16" title="Simao, F. A., Waterhouse, R. M., Ioannidis, P., Kriventseva, E. V. & Zdobnov, E. M. BUSCO: assessing genome assembly and annotation completeness with single-copy orthologs. Bioinformatics 31, 3210–3212 (2015)." href="/articles/s41477-019-0588-4#ref-CR16" id="ref-link-section-d326961673e1154">16</a></sup> (85.04% complete gene models plus 4.60% fragmented gene models), compared with 93.51% and 92.15% captured in <i>P. patens</i><sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 14" title="Rensing, S. A. et al. The Physcomitrella genome reveals evolutionary insights into the conquest of land by plants. Science 319, 64–69 (2008)." href="/articles/s41477-019-0588-4#ref-CR14" id="ref-link-section-d326961673e1161">14</a></sup> and <i>M. polymorpha</i><sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 15" title="Bowman, J. L. et al. Insights into land plant evolution garnered from the Marchantia polymorpha genome. Cell 171, 287–304 (2017)." href="/articles/s41477-019-0588-4#ref-CR15" id="ref-link-section-d326961673e1167">15</a></sup>, respectively (Supplementary Table <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">10</a>). In addition to protein-coding genes, we also identified 30 known mature micro RNAs (miRNAs), 180 novel mature miRNAs, 347 transfer RNAs, 94 ribosomal RNAs and 83 small nuclear RNAs (snRNAs) in the <i>A. angustus</i> genome (Supplementary Table <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">11</a>). Nine mature miRNA sequences that appear conserved among land plants (miR156/157, miR159/319, miR160, miR165/166, miR170/171, miR408, miR477, miR535 and miR536)<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 17" title="Axtell, M. J. & Bowman, J. L. Evolution of plant microRNAs and their targets. Trends Plant Sci. 13, 343–349 (2008)." href="/articles/s41477-019-0588-4#ref-CR17" id="ref-link-section-d326961673e1181">17</a></sup> were also found in <i>A. angustus</i> (Supplementary Table <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">12</a>).</p><div class="c-article-table" data-test="inline-table" data-container-section="table" id="table-1"><figure><figcaption class="c-article-table__figcaption"><b id="Tab1" data-test="table-caption">Table 1 Assembly and annotation statistics of the draft genome of <i>A. angustus</i></b></figcaption><div class="u-text-right u-hide-print"><a class="c-article__pill-button" data-test="table-link" data-track="click" data-track-action="view table" data-track-label="button" rel="nofollow" href="/articles/s41477-019-0588-4/tables/1" aria-label="Full size table 1"><span>Full size table</span><svg width="16" height="16" focusable="false" role="img" aria-hidden="true" class="u-icon"><use xmlns:xlink="http://www.w3.org/1999/xlink" xlink:href="#icon-eds-i-chevron-right-small"></use></svg></a></div></figure></div></div></div></section><section data-title="Comparative genomic analysis"><div class="c-article-section" id="Sec3-section"><h2 class="c-article-section__title js-section-title js-c-reading-companion-sections-item" id="Sec3">Comparative genomic analysis</h2><div class="c-article-section__content" id="Sec3-content"><p>For sequence similarity-based clustering of homologues, we used the predicted proteomes of <i>A. angustus</i> and 18 other green plants with fully-sequenced genomes (that is, 11 other land plants, two charophyte green algae and five chlorophyte green algae; Supplementary Table <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">13</a>). Genes of <i>A. angustus</i> are distributed among 7,644 gene families that are shared with other plants, and 497 gene families that appear to be unique to <i>A. angustus</i> (Fig. <a data-track="click" data-track-label="link" data-track-action="figure anchor" href="/articles/s41477-019-0588-4#Fig1">1a</a> and Supplementary Table <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">14</a>). In the shared gene families, most <i>A. angustus</i> genes (that is, 9,680) cluster with land plant genes, and only a very small number (that is, 107) specifically cluster with green algae genes (Supplementary Fig. <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">7</a>). The gene families unique to <i>A. angustus</i> are enriched in various biosynthetic categories (for example, terpenoid and zeatin) and various activity categories (for example, nutrient reservoir activity and catechol oxidase activity) (Supplementary Tables <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">15</a> and <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">16</a>).</p><div class="c-article-section__figure js-c-reading-companion-figures-item" data-test="figure" data-container-section="figure" id="figure-1" data-title="Comparative genomic analysis of A. angustus and 18 other plant species."><figure><figcaption><b id="Fig1" class="c-article-section__figure-caption" data-test="figure-caption-text">Fig. 1: Comparative genomic analysis of <i>A. angustus</i> and 18 other plant species.</b></figcaption><div class="c-article-section__figure-content"><div class="c-article-section__figure-item"><a class="c-article-section__figure-link" data-test="img-link" data-track="click" data-track-label="image" data-track-action="view figure" href="/articles/s41477-019-0588-4/figures/1" rel="nofollow"><picture><source type="image/webp" srcset="//media.springernature.com/lw685/springer-static/image/art%3A10.1038%2Fs41477-019-0588-4/MediaObjects/41477_2019_588_Fig1_HTML.png?as=webp"><img aria-describedby="Fig1" src="//media.springernature.com/lw685/springer-static/image/art%3A10.1038%2Fs41477-019-0588-4/MediaObjects/41477_2019_588_Fig1_HTML.png" alt="figure 1" loading="lazy" width="685" height="725"></picture></a></div><div class="c-article-section__figure-description" data-test="bottom-caption" id="figure-1-desc"><p><b>a</b>, Comparison of the number of gene families identified by OrthoMCL. The Venn diagram shows the shared and unique gene families in <i>A. angustus</i>, Setaphyta, Tracheophyta, Charophyta and Chlorophyta. The gene-family number is listed in each of the components. <b>b</b>, Gene-family gain (+)/loss (−) among 19 green plants. The numbers of gained (blue) and lost (red) gene families are shown above the branches. The boxed number indicates the gene-family size at each node. The number of gene families, orphans (single-copy gene families) and number of predicted genes is indicated next to each species. <b>c</b>, Comparison of whole paranome, anchor pair and one-to-one orthologue distribution of the number of synonymous substitutions per synonymous site (<i>K</i><sub>S</sub>) across the three bryophyte species (<i>P. patens</i>, <i>M. polymorpha</i> and <i>A. angustus</i>).</p></div></div><div class="u-text-right u-hide-print"><a class="c-article__pill-button" data-test="article-link" data-track="click" data-track-label="button" data-track-action="view figure" href="/articles/s41477-019-0588-4/figures/1" data-track-dest="link:Figure1 Full size image" aria-label="Full size image figure 1" rel="nofollow"><span>Full size image</span><svg width="16" height="16" focusable="false" role="img" aria-hidden="true" class="u-icon"><use xmlns:xlink="http://www.w3.org/1999/xlink" xlink:href="#icon-eds-i-chevron-right-small"></use></svg></a></div></figure></div><p>Phylogenetic inferences from 85 single-copy nuclear genes sampled for <i>A. angustus</i> and 18 other green plants resolve hornworts (<i>A. angustus</i>), mosses (<i>P. patens</i>) and liverworts (<i>M. polymorpha</i>) as a monophyletic group, with hornworts sister to mosses and liverworts, which agrees with inferences from 852 nuclear genes sampled from 103 plant species<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 1" title="Puttick, M. N. et al. The interrelationships of land plants and the nature of the ancestral embryophyte. Curr. Biol. 28, 733–745 (2018)." href="/articles/s41477-019-0588-4#ref-CR1" id="ref-link-section-d326961673e1532">1</a></sup> (Fig. <a data-track="click" data-track-label="link" data-track-action="figure anchor" href="/articles/s41477-019-0588-4#Fig1">1b</a>, Supplementary Figs. <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">8</a> and <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">9</a>, Supplementary Table <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">17</a> and Supplementary Notes <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">2.1</a> and <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">2.2</a>). The divergence (Supplementary Figs. <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">10</a> and <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">11</a>, Supplementary Tables <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">18</a>–<a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">20</a> and Supplementary Note <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">S2.3</a>) of the extant crown group of hornworts is estimated at 275.62 million years ago (Ma) (95% highest posterior density, 179.3–384.6 Ma) (middle Carboniferous–early Jurassic) (Supplementary Fig. <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">11</a> and Supplementary Table <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">20</a>), which is comparable to the crown age of hornworts estimated based on two organellar sequences from 77 hornworts and 11 other land plants<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 10" title="Villarreal, J. C. & Renner, S. S. Hornwort pyrenoids, carbon-concentrating structures, evolved and were lost at least five times during the last 100 million years. Proc. Natl Acad. Sci. USA 109, 18873–18878 (2012)." href="/articles/s41477-019-0588-4#ref-CR10" id="ref-link-section-d326961673e1578">10</a></sup>. These estimates are thus older than those inferred from the fossil record, considering that the oldest putative hornwort fossil is a spore from the Lower Cretaceous Baqueró Formation, Argentina (from 145 to 100 Ma) that resembles the spores of extant <i>Anthoceros</i><sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 18" title="Archangelsky, S. & Villar de Seone, L. Estudios palinógicos de la formación Baqueró (Cretácico), provincia de Santa Cruz, Argentina. Ameghiniana 35, 7–19 (1996)." href="/articles/s41477-019-0588-4#ref-CR18" id="ref-link-section-d326961673e1584">18</a></sup>.</p><p>Comparative genomics shows that the genome of <i>A. angustus</i> has lost many gene families (that is, 2,145) and comparatively only modest gains (that is, 497) (Fig. <a data-track="click" data-track-label="link" data-track-action="figure anchor" href="/articles/s41477-019-0588-4#Fig1">1b</a>). A similar trend characterizes the genome of <i>Marchantia</i> and of the ancestor common to all bryophytes, whereas <i>P. patens</i> has gained more families (that is, 1,334) than it has lost (that is, 1,248; Fig. <a data-track="click" data-track-label="link" data-track-action="figure anchor" href="/articles/s41477-019-0588-4#Fig1">1b</a>). Thus, bryophyte genomes may not only harbour a number of genes and gene families comparable to those of vascular plants and in particular seed plants (Fig. <a data-track="click" data-track-label="link" data-track-action="figure anchor" href="/articles/s41477-019-0588-4#Fig1">1b</a>) but may also be highly dynamic through evolutionary time.</p><p>Many, if not most, land plants harbour genomic signatures of ancient whole-genome duplication (WGD)<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 19" title="Van de Peer, Y., Mizrachi, E. & Marchal, K. The evolutionary significance of polyploidy. Nat. Rev. Genet. 18, 411–424 (2017)." href="/articles/s41477-019-0588-4#ref-CR19" id="ref-link-section-d326961673e1613">19</a></sup>. However, like that of <i>Marchantia</i><sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 15" title="Bowman, J. L. et al. Insights into land plant evolution garnered from the Marchantia polymorpha genome. Cell 171, 287–304 (2017)." href="/articles/s41477-019-0588-4#ref-CR15" id="ref-link-section-d326961673e1619">15</a></sup>, the genome of <i>Anthoceros</i> lacks evidence of having undergone a WGD (Fig. <a data-track="click" data-track-label="link" data-track-action="figure anchor" href="/articles/s41477-019-0588-4#Fig1">1c</a>, Supplementary Fig. <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">13</a> and Supplementary Note <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">3.1</a>), which confirms the hypothesis drawn previously from the analysis of transcriptomic data<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 20" title="Lang, D. et al. The Physcomitrella patens chromosome-scale assembly reveals moss genome structure and evolution. Plant J. 93, 515–533 (2018)." href="/articles/s41477-019-0588-4#ref-CR20" id="ref-link-section-d326961673e1636">20</a></sup>. The chromosomal arrangement of genes is not much conserved among the three bryophyte lineages (Supplementary Fig. <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">14a,b</a> and Supplementary Note <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">3.2</a>), which likely reflects the ancient divergence of these different lineages of bryophytes. For example, the longest co-linear block corresponds to a mere five anchor pairs for both <i>A. angustus</i> versus <i>P. patens</i> and <i>A. angustus</i> versus <i>M. polymorpha</i>, whereas within the <i>A. angustus</i> genome, the largest co-linear segment consists of six anchor pairs (Supplementary Fig. <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">14</a>).</p><p>The <i>A. angustus</i> genome contains a much lower percentage of multi-copy gene families than that of single-copy gene families, implying low genetic redundancy (Supplementary Table <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">17</a>), which is similar to what has been observed for the liverwort <i>Marchantia</i><sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 15" title="Bowman, J. L. et al. Insights into land plant evolution garnered from the Marchantia polymorpha genome. Cell 171, 287–304 (2017)." href="/articles/s41477-019-0588-4#ref-CR15" id="ref-link-section-d326961673e1677">15</a></sup>.</p></div></div></section><section data-title="Transcription factors"><div class="c-article-section" id="Sec4-section"><h2 class="c-article-section__title js-section-title js-c-reading-companion-sections-item" id="Sec4">Transcription factors</h2><div class="c-article-section__content" id="Sec4-content"><p>The <i>A. angustus</i> genome comprises 333 putative transcription factor (TF) genes covering 61 families, a number that is highly similar to that of the other two bryophyte genomes (Supplementary Fig. <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">15</a>, Supplementary Table <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM3">21</a> and Supplementary Note <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">4.1</a>). The diversity of TF genes in extant plants is rather stable (Supplementary Fig. <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">15</a>) and resulted from two ancient bursts of TF families during the diversification of green plants: one concomitant with the origin of streptophytes and the other with the transition to land<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 15" title="Bowman, J. L. et al. Insights into land plant evolution garnered from the Marchantia polymorpha genome. Cell 171, 287–304 (2017)." href="/articles/s41477-019-0588-4#ref-CR15" id="ref-link-section-d326961673e1705">15</a>,<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 21" title="Catarino, B., Hetherington, A. J., Emms, D. M., Kelly, S. & Dolan, L. The stepwise increase in the number of transcription factor families in the Precambrian predated the diversification of plants on land. Mol. Biol. Evol. 33, 2815–2819 (2016)." href="/articles/s41477-019-0588-4#ref-CR21" id="ref-link-section-d326961673e1708">21</a></sup>. In plants, genes encoding TFs are among the most highly retained following polyploidy<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 22" title="Cheng, F. et al. Gene retention, fractionation and subgenome differences in polyploid plants. Nat. Plants 4, 258–268 (2018)." href="/articles/s41477-019-0588-4#ref-CR22" id="ref-link-section-d326961673e1712">22</a></sup>, a pattern reflected in the comparison of the three bryophyte genomes<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 14" title="Rensing, S. A. et al. The Physcomitrella genome reveals evolutionary insights into the conquest of land by plants. Science 319, 64–69 (2008)." href="/articles/s41477-019-0588-4#ref-CR14" id="ref-link-section-d326961673e1716">14</a>,<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 15" title="Bowman, J. L. et al. Insights into land plant evolution garnered from the Marchantia polymorpha genome. Cell 171, 287–304 (2017)." href="/articles/s41477-019-0588-4#ref-CR15" id="ref-link-section-d326961673e1719">15</a></sup>. <i>A. angustus</i> and <i>M. polymorpha</i>, whose genome did not undergo WGDs hold a small number of TF compared to <i>P. patens</i>, which experienced at least one WGD in its ancestry, resulting in a substantially larger number of TF genes (Supplementary Fig. <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">15</a>). It supports the hypothesis that the WGD is an important mechanism for expansion of TF families<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 23" title="Lang, D. et al. Genome-wide phylogenetic comparative analysis of plant transcriptional regulation: a timeline of loss, gain, expansion, and correlation with complexity. Genome Biol. Evol. 2, 488–503 (2010)." href="/articles/s41477-019-0588-4#ref-CR23" id="ref-link-section-d326961673e1736">23</a></sup>.</p><p>Phylogenetic analyses of 24 gene families contributing to the development of plant body plans or adaptation to the terrestrial environment, including 16 TF gene families<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 24" title="Sakakibara, K. Technological innovations give rise to a new era of plant evolutionary developmental biology. Adv. Bot. Res. 78, 3–35 (2016)." href="/articles/s41477-019-0588-4#ref-CR24" id="ref-link-section-d326961673e1743">24</a>,<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 25" title="Szövényi, P., Waller, M. & Kirbis, A. Evolution of the plant body plan. Curr. Top. Dev. Biol. 131, 1–34 (2019)." href="/articles/s41477-019-0588-4#ref-CR25" id="ref-link-section-d326961673e1746">25</a></sup> (Fig. <a data-track="click" data-track-label="link" data-track-action="figure anchor" href="/articles/s41477-019-0588-4#Fig2">2a</a>, Supplementary Figs. <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">16</a>–<a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">54</a>, Supplementary Table <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM3">22</a> and Supplementary Note <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">4.2</a>), confirm that a considerable number of genes, such as genes involved in gametophyte or sporophyte development, haploid–diploid transition, meristem development, filamentous growth, photomorphogenesis and auxin signalling (Fig. <a data-track="click" data-track-label="link" data-track-action="figure anchor" href="/articles/s41477-019-0588-4#Fig2">2</a>), composed the genetic toolkit of plants before the conquest of land<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 26" title="Floyd, S. K. & Bowman, J. L. The ancestral developmental tool kit of land plants. Int. J. Plant Sci. 168, 1–35 (2007)." href="/articles/s41477-019-0588-4#ref-CR26" id="ref-link-section-d326961673e1769">26</a></sup>. In particular, the TF genes for filamentous growth and auxin signalling arose in charophyte green algae<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 27" title="Hori, K. et al. Klebsormidium flaccidum genome reveals primary factors for plant terrestrial adaptation. Nat. Commun. 5, 3978 (2014)." href="/articles/s41477-019-0588-4#ref-CR27" id="ref-link-section-d326961673e1773">27</a>,<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 28" title="Nishiyama, T. et al. The Chara genome: secondary complexity and implications for plant terrestrialization. Cell 174, 448–464 (2018)." href="/articles/s41477-019-0588-4#ref-CR28" id="ref-link-section-d326961673e1776">28</a></sup> (Fig. <a data-track="click" data-track-label="link" data-track-action="figure anchor" href="/articles/s41477-019-0588-4#Fig2">2b</a>), which are thought to be the closest living relatives to extant land plants, implying the preliminary establishment of relatively more complex body plan in these basal streptophytes for plant terrestrial adaptation<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 29" title="Wodniok, S. et al. Origin of land plants: do conjugating green algae hold the key? BMC Evol. Biol. 11, 104 (2011)." href="/articles/s41477-019-0588-4#ref-CR29" id="ref-link-section-d326961673e1783">29</a></sup>. Furthermore, a set of genes underlying key morphological innovations for terrestrial adaptation probably evolved along with the colonization of land<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 30" title="Ishizaki, K. Evolution of land plants: insights from molecular studies on basal lineages. Biosci. Biotechnol. Biochem. 81, 73–80 (2017)." href="/articles/s41477-019-0588-4#ref-CR30" id="ref-link-section-d326961673e1788">30</a>,<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 31" title="Rensing, S. A. Great moments in evolution: the conquest of land by plants. Curr. Opin. Plant Biol. 42, 49–54 (2018)." href="/articles/s41477-019-0588-4#ref-CR31" id="ref-link-section-d326961673e1791">31</a></sup> (Fig. <a data-track="click" data-track-label="link" data-track-action="figure anchor" href="/articles/s41477-019-0588-4#Fig2">2b</a>), such as <i>SMF</i> and <i>ICE</i> for stomatal development (Supplementary Figs. <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">29</a> and <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">30</a>), <i>APB</i>, <i>CLE</i> and <i>CLV1</i> for 3D growth (Supplementary Figs. <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">36</a> and <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">50</a>–<a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">52</a>), and <i>VNS</i> for water-conducting-cell development (Supplementary Fig. <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">38</a>). The sporophyte morphology of bryophytes is relatively simple, and many of the genes involved in the elaborate regulation of embryogenesis<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 32" title="Braybrook, S. A. & Harada, J. J. LECs go crazy in embryo development. Trends Plant Sci. 13, 624–630 (2008)." href="/articles/s41477-019-0588-4#ref-CR32" id="ref-link-section-d326961673e1836">32</a></sup>, such as <i>FUS3</i>, <i>LEC1</i>, <i>LEC2</i>, <i>NF-YA1/9</i> and <i>NF-YA3/5/6/8</i> are absent in <i>A. angustus</i>, <i>Marchantia</i> and <i>Physcomitrella</i> (Fig. <a data-track="click" data-track-label="link" data-track-action="figure anchor" href="/articles/s41477-019-0588-4#Fig2">2a</a> and Supplementary Figs. <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">39</a>–<a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">41</a>). The <i>ABI3</i> genes that mainly function in embryo maturation and seed desiccation tolerance in flowering plants are present in bryophytes, and have roles in desiccation tolerance in their vegetative tissues<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 33" title="Takezawa, D., Komatsu, K. & Sakata, Y. ABA in bryophytes: how a universal growth regulator in life became a plant hormone? J. Plant Res. 124, 437–453 (2011)." href="/articles/s41477-019-0588-4#ref-CR33" id="ref-link-section-d326961673e1878">33</a></sup>.</p><div class="c-article-section__figure js-c-reading-companion-figures-item" data-test="figure" data-container-section="figure" id="figure-2" data-title="Major TFs for plant body plan and evolutionary innovations within plants."><figure><figcaption><b id="Fig2" class="c-article-section__figure-caption" data-test="figure-caption-text">Fig. 2: Major TFs for plant body plan and evolutionary innovations within plants.</b></figcaption><div class="c-article-section__figure-content"><div class="c-article-section__figure-item"><a class="c-article-section__figure-link" data-test="img-link" data-track="click" data-track-label="image" data-track-action="view figure" href="/articles/s41477-019-0588-4/figures/2" rel="nofollow"><picture><source type="image/webp" srcset="//media.springernature.com/lw685/springer-static/image/art%3A10.1038%2Fs41477-019-0588-4/MediaObjects/41477_2019_588_Fig2_HTML.png?as=webp"><img aria-describedby="Fig2" src="//media.springernature.com/lw685/springer-static/image/art%3A10.1038%2Fs41477-019-0588-4/MediaObjects/41477_2019_588_Fig2_HTML.png" alt="figure 2" loading="lazy" width="685" height="1096"></picture></a></div><div class="c-article-section__figure-description" data-test="bottom-caption" id="figure-2-desc"><p><b>a</b>, Overview for the number of major TFs for plant body plan in ten green plants. Colour key on the upper left of the heatmap denotes the TF numbers. <b>b</b>, Major innovations in plants and evolutionary features of three bryophyte lineages.</p></div></div><div class="u-text-right u-hide-print"><a class="c-article__pill-button" data-test="article-link" data-track="click" data-track-label="button" data-track-action="view figure" href="/articles/s41477-019-0588-4/figures/2" data-track-dest="link:Figure2 Full size image" aria-label="Full size image figure 2" rel="nofollow"><span>Full size image</span><svg width="16" height="16" focusable="false" role="img" aria-hidden="true" class="u-icon"><use xmlns:xlink="http://www.w3.org/1999/xlink" xlink:href="#icon-eds-i-chevron-right-small"></use></svg></a></div></figure></div><p>In <i>A. angustus</i>, most genes involved in the development of plant body plans have a single copy, and a few <i>A. angustus</i> TF gene families even lost a subset of duplicates (Fig. <a data-track="click" data-track-label="link" data-track-action="figure anchor" href="/articles/s41477-019-0588-4#Fig2">2a</a> and Supplementary Figs. <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">16</a>–<a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">52</a>). For example, in the bHLH family, the class I <i>RSL</i> gene that controls the development of rhizoids and root hairs, thought to have been important for the colonization of land<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 34" title="Proust, H. et al. RSL class I genes controlled the development of epidermal structures in the common ancestor of land plants. Curr. Biol. 26, 93–99 (2016)." href="/articles/s41477-019-0588-4#ref-CR34" id="ref-link-section-d326961673e1929">34</a></sup>, is present in the <i>A. angustus</i> genome, whereas the class II <i>RSL</i> genes responsible for regulating protonema differentiation in <i>P. patens</i> or root hair elongation in <i>A. thaliana</i> by auxin<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 35" title="Pires, N. D. et al. Recruitment and remodeling of an ancient gene regulatory network during land plant evolution. Proc. Natl Acad. Sci. USA 110, 9571–9576 (2013)." href="/articles/s41477-019-0588-4#ref-CR35" id="ref-link-section-d326961673e1946">35</a></sup> are absent (Supplementary Fig. <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">27</a> and Supplementary Note <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">4.2</a>). The lack of class II <i>RSL</i> genes in <i>A. angustus</i> might be related to the morphological simplification of this species with respect to tip-growing filamentous structures<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2" title="Goffinet, B. & Buck, W. R. in The Evolution of Plant Form (eds Ambrose, B. & Purruganan, M.) 51–90 (Wiley–Blackwell, 2013)." href="/articles/s41477-019-0588-4#ref-CR2" id="ref-link-section-d326961673e1962">2</a></sup>. For the <i>KNOX</i> genes from the homeobox gene family, the <i>A. angustus</i> genome retains one class II <i>KNOX</i> gene for haploid-to-diploid morphological transition<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 36" title="Sakakibara, K. et al. KNOX2 genes regulate the haploid-to-diploid morphological transition in land plants. Science 339, 1067–1070 (2013)." href="/articles/s41477-019-0588-4#ref-CR36" id="ref-link-section-d326961673e1976">36</a></sup>, but lacks class I <i>KNOX</i> genes (Supplementary Fig. <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">23</a>), whose activity is necessary for seta extension in the sporophytes in <i>P. patens</i><sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 37" title="Coudert, Y., Novák, O. & Harrison, C. J. A KNOX-cytokinin regulatory module predates the origin of indeterminate vascular plants. Curr. Biol. 29, 2743–2750 (2019)." href="/articles/s41477-019-0588-4#ref-CR37" id="ref-link-section-d326961673e1989">37</a></sup>. The absence of this gene might be linked to the absence of setae in hornworts<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2" title="Goffinet, B. & Buck, W. R. in The Evolution of Plant Form (eds Ambrose, B. & Purruganan, M.) 51–90 (Wiley–Blackwell, 2013)." href="/articles/s41477-019-0588-4#ref-CR2" id="ref-link-section-d326961673e1993">2</a></sup>. The genome of <i>A. angustus</i> also holds few type II MIKC<sup>C</sup> <i>MADS-box</i>, class B <i>ARF</i>, <i>NCARF</i> and short <i>PIN</i> genes, as a result of gene losses suggested by our phylogenetic analysis (Supplementary Figs. <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">17</a>, <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">42</a>, <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">45</a> and Supplementary Note <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">4.2</a>). The class II <i>RSL</i>, class B <i>ARF</i>, <i>NCARF</i> and short <i>PIN</i> genes all have auxin-related functions (Supplementary Note <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">4.2</a>). Since these auxin-related genes were consistently lost in <i>A. angustus</i>, this hornwort species possesses the simplest auxin molecular toolkit among all investigated land plants so far<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 38" title="Mutte, S. K. et al. Origin and evolution of the nuclear auxin response system. eLife 7, e33399 (2018)." href="/articles/s41477-019-0588-4#ref-CR38" id="ref-link-section-d326961673e2047">38</a></sup>. Thus, like the liverwort <i>M. polymorpha</i><sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 15" title="Bowman, J. L. et al. Insights into land plant evolution garnered from the Marchantia polymorpha genome. Cell 171, 287–304 (2017)." href="/articles/s41477-019-0588-4#ref-CR15" id="ref-link-section-d326961673e2053">15</a></sup>, <i>A. angustus</i> exhibits low redundancy for genes shaping the plant body plan (Fig. <a data-track="click" data-track-label="link" data-track-action="figure anchor" href="/articles/s41477-019-0588-4#Fig2">2b</a>). Such a limited toolkit may be characteristic of the ancestor to bryophytes and hence, perhaps, of the earliest land plants with a dominant thalloid gametophyte, and provide the foundation to explaining the architectural simplicity of these plants. By contrast, the genome of <i>P. patens</i>, which develops a leafy stem, has the most TF genes involved in the development of plant body plans among the compared bryophytes (Fig. <a data-track="click" data-track-label="link" data-track-action="figure anchor" href="/articles/s41477-019-0588-4#Fig2">2b</a>). Although the genome of <i>A. angustus</i> seems poor in genes composing the network underlying the development of its body plan, the TF gene families linked to responses to terrestrial environmental stimuli exhibit lineage-specific gene expansions in <i>A. angustus</i>, namely, the <i>LISCL</i> genes for mycorrhizal signalling in the <i>GRAS</i> gene family<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 39" title="Cenci, A. & Rouard, M. Evolutionary analyses of GRAS transcription factors in angiosperms. Front. Plant Sci. 8, 273 (2017)." href="/articles/s41477-019-0588-4#ref-CR39" id="ref-link-section-d326961673e2083">39</a></sup> (Supplementary Fig. <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">53</a>) and the clade <i>SIP1</i> for ABA signalling under water stress in the <i>Trihelix</i> gene family<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 40" title="Kaplan-Levy, R. N., Brewer, P. B., Quon, T. & Smyth, D. R. The trihelix family of transcription factors—light, stress and development. Trends Plant Sci. 17, 163–171 (2012)." href="/articles/s41477-019-0588-4#ref-CR40" id="ref-link-section-d326961673e2096">40</a></sup> (Supplementary Fig. <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">54</a>).</p></div></div></section><section data-title="Gene-family expansion"><div class="c-article-section" id="Sec5-section"><h2 class="c-article-section__title js-section-title js-c-reading-companion-sections-item" id="Sec5">Gene-family expansion</h2><div class="c-article-section__content" id="Sec5-content"><p>Besides two TF gene families, the <i>A. angustus</i> genome harbours a variety of other uniquely expanded gene families (Supplementary Fig. <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">55</a>). The genome comprises an very large number of pentatricopeptide repeat (<i>PPR</i>) genes for plant organellar RNA processing<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 41" title="Fujii, S. & Small, I. The evolution of RNA editing and pentatricopeptide repeat genes. N. Phytol. 191, 37–47 (2011)." href="/articles/s41477-019-0588-4#ref-CR41" id="ref-link-section-d326961673e2120">41</a></sup>, accounting for approximately 7.90% of the predicted protein-coding genes. The expanded <i>PPR</i> genes are PLS-class <i>PPR</i> genes (Supplementary Fig. <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">55</a>, Supplementary Tables <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM3">23</a> and <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM3">24</a> and Supplementary Note <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">5.1</a>). Most of the PLS-class PPR proteins in <i>A. angustus</i> were predicted to be localized in the mitochondrion or chloroplast (Supplementary Table <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM3">24</a>). The expansion of the PLS-class <i>PPR</i> genes correlates with the large number of RNA editing sites estimated in the organellar genomes of <i>A. angustus</i> (Supplementary Table <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM3">23</a>). Our findings add further support to the hypothesis that an increase in the number of both RNA editing sites and <i>PPR</i> genes (especially the PLS-class PPR) occurred after the separation of land plants from green algae<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 41" title="Fujii, S. & Small, I. The evolution of RNA editing and pentatricopeptide repeat genes. N. Phytol. 191, 37–47 (2011)." href="/articles/s41477-019-0588-4#ref-CR41" id="ref-link-section-d326961673e2162">41</a>,<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 42" title="Cheng, S. et al. Redefining the structural motifs that determine RNA binding and RNA editing by pentatricopeptide repeat proteins in land plants. Plant J. 85, 532–547 (2016)." href="/articles/s41477-019-0588-4#ref-CR42" id="ref-link-section-d326961673e2165">42</a></sup> (Supplementary Table <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM3">23</a>). The reduced number of <i>PPR</i> genes and absence of RNA editing in marchantiid liverworts are most probably secondary losses (Supplementary Table <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM3">23</a>), as the organellar RNA editing and plant-specific extensions of <i>PPR</i> genes were also found in jungermanniid liverworts<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 43" title="Rüdinger, M., Polsakiewicz, M. & Knoop, V. Organellar RNA editing and plant-specific extensions of pentatricopeptide repeat proteins in jungermanniid but not in marchantiid liverworts. Mol. Biol. Evol. 25, 1405–1414 (2008)." href="/articles/s41477-019-0588-4#ref-CR43" id="ref-link-section-d326961673e2182">43</a></sup>. Through RNA editing, the PPR proteins could act as ‘repair’ factors that alleviate DNA damage caused by increased UV exposure in terrestrial environments<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 41" title="Fujii, S. & Small, I. The evolution of RNA editing and pentatricopeptide repeat genes. N. Phytol. 191, 37–47 (2011)." href="/articles/s41477-019-0588-4#ref-CR41" id="ref-link-section-d326961673e2186">41</a></sup>. Other stress-response gene families have also expanded in <i>A. angustus</i>, such as cupin and cytochrome P450 (CYP) (Supplementary Fig. <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">55</a>). Two groups of cupin (PF00190) proteins—that is, monocupins and bicupins—can be recognized on the basis of the number of cupin domains<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 44" title="Dunwell, J. M., Khuri, S. & Gane, P. J. Microbial relatives of the seed storage proteins of higher plants: conservation of structure and diversification of function during evolution of the cupin superfamily. Microbiol. Mol. Biol. Rev. 64, 153–179 (2000)." href="/articles/s41477-019-0588-4#ref-CR44" id="ref-link-section-d326961673e2197">44</a></sup>. In <i>A. angustus</i>, the <i>cupin</i> gene family has undergone a significant expansion (Supplementary Table <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM3">25</a>) such that it comprises more <i>bicupin</i> genes than any other plant (Fig. <a data-track="click" data-track-label="link" data-track-action="figure anchor" href="/articles/s41477-019-0588-4#Fig3">3a</a>, Supplementary Figs. <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">56</a> and <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">57</a>, Supplementary Table <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM3">25</a> and Supplementary Note <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">5.2</a>). Expansion of the <i>cupin</i> gene family in <i>A. angustus</i> resulted mainly from tandem gene duplications (Fig. <a data-track="click" data-track-label="link" data-track-action="figure anchor" href="/articles/s41477-019-0588-4#Fig3">3b,c</a> and Supplementary Note <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">5.2</a>). Since bicupins (that is, 11S and 7S seed storage proteins) are desiccation-tolerant proteins in higher land plants<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 44" title="Dunwell, J. M., Khuri, S. & Gane, P. J. Microbial relatives of the seed storage proteins of higher plants: conservation of structure and diversification of function during evolution of the cupin superfamily. Microbiol. Mol. Biol. Rev. 64, 153–179 (2000)." href="/articles/s41477-019-0588-4#ref-CR44" id="ref-link-section-d326961673e2242">44</a></sup>, the large number of <i>bicupin</i> genes in <i>A. angustus</i> could indicate adaptation for coping with drought stress in the terrestrial environment. The large number of <i>A. angustus</i>-specific <i>monocupin</i> genes are homologous to the <i>P. patens PpGLP6</i> gene (XP_001782709.1) (Supplementary Fig. <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">57</a> and Supplementary Note <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">5.2</a>), which encodes a protein with manganese-containing extracellular superoxide dismutase (SOD) activity to respond to oxidative stress in terrestrial environments<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 45" title="Nakata, M. et al. Germin-like protein gene family of a moss, Physcomitrella patens, phylogenetically falls into two characteristic new clades. Plant Mol. Biol. 56, 381–395 (2004)." href="/articles/s41477-019-0588-4#ref-CR45" id="ref-link-section-d326961673e2269">45</a></sup>. The <i>CYP</i> genes for primary and secondary metabolism have also expanded in <i>A. angustus</i> (Supplementary Fig. <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">55</a> and Supplementary Note <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">5.3</a>). For instance, genes belonging to the subfamilies <i>CYP71</i> and <i>CYP85</i> contain 56 and 46 genes, respectively (Supplementary Figs. <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">58</a>–<a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">61</a> and Supplementary Tables <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM3">26</a> and <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM3">27</a>). The <i>A. angustus CYP</i> genes were assigned to 28 KEGG pathways, of which ‘flavonoid 3′-monooxygenase/flavonoid 3′,5′-hydroxylase’ and ‘abscisic acid 8′-hydroxylase’ were the most representative (Supplementary Table <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM3">28</a>). Within the <i>CYP71</i> gene subfamily, genes homologous to flavonoid 3′-hydroxylase (monooxygenase) (<i>F3'H</i>) or flavonoid 3′,5′-hydroxylase (<i>F3</i>′<i>5</i>′<i>H</i>) genes that are involved in flavonoid biosynthesis<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 46" title="Pollastri, S. & Tattini, M. Flavonols: old compounds for old roles. Ann. Bot. 108, 1225–1233 (2011)." href="/articles/s41477-019-0588-4#ref-CR46" id="ref-link-section-d326961673e2327">46</a></sup> are highly expanded in <i>A. angustus</i> (Supplementary Fig. <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">59</a> and Supplementary Note <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">5.3</a>). Because flavonoids have an important role in UV-B protection<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 46" title="Pollastri, S. & Tattini, M. Flavonols: old compounds for old roles. Ann. Bot. 108, 1225–1233 (2011)." href="/articles/s41477-019-0588-4#ref-CR46" id="ref-link-section-d326961673e2340">46</a></sup>, the expansion of flavonoid biosynthesis related genes in <i>A. angustus</i> might again represent a molecular adaptation to life in the terrestrial environment. Among the <i>CYP85</i> genes, the genes homologous to abscisic acid 8′-hydroxylase genes involved in abscisic acid catabolism during drought stress response<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 47" title="Sakata, Y., Komatsu, K. & Takezawa, D. in Progress in Botany (ed. Lüttge, U.) 57–96 (Springer-Verlag, 2014)." href="/articles/s41477-019-0588-4#ref-CR47" id="ref-link-section-d326961673e2351">47</a></sup> are also uniquely abundant in <i>A. angustus</i> (Supplementary Fig. <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">60</a> and Supplementary Note <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">5.3</a>), and may account for the high desiccation tolerance of <i>A. angustus</i>. Like the <i>cupin</i> gene family, many of the above expanded gene families occur in tandem arrays (Supplementary Table <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM3">29</a>). At least 9.82% of protein-coding genes in <i>A. angustus</i> form ‘tandem’ clusters in the genome (Supplementary Table <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM3">30</a> and Supplementary Note <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">5.4</a>), compared with only 1% in <i>P. patens</i><sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 14" title="Rensing, S. A. et al. The Physcomitrella genome reveals evolutionary insights into the conquest of land by plants. Science 319, 64–69 (2008)." href="/articles/s41477-019-0588-4#ref-CR14" id="ref-link-section-d326961673e2386">14</a></sup> and 5.9% in <i>M. polymorpha</i><sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 15" title="Bowman, J. L. et al. Insights into land plant evolution garnered from the Marchantia polymorpha genome. Cell 171, 287–304 (2017)." href="/articles/s41477-019-0588-4#ref-CR15" id="ref-link-section-d326961673e2392">15</a></sup>.</p><div class="c-article-section__figure js-c-reading-companion-figures-item" data-test="figure" data-container-section="figure" id="figure-3" data-title="Expansion of cupin gene family in A. angustus."><figure><figcaption><b id="Fig3" class="c-article-section__figure-caption" data-test="figure-caption-text">Fig. 3: Expansion of <i>cupin</i> gene family in <i>A. angustus</i>.</b></figcaption><div class="c-article-section__figure-content"><div class="c-article-section__figure-item"><a class="c-article-section__figure-link" data-test="img-link" data-track="click" data-track-label="image" data-track-action="view figure" href="/articles/s41477-019-0588-4/figures/3" rel="nofollow"><picture><source type="image/webp" srcset="//media.springernature.com/lw685/springer-static/image/art%3A10.1038%2Fs41477-019-0588-4/MediaObjects/41477_2019_588_Fig3_HTML.png?as=webp"><img aria-describedby="Fig3" src="//media.springernature.com/lw685/springer-static/image/art%3A10.1038%2Fs41477-019-0588-4/MediaObjects/41477_2019_588_Fig3_HTML.png" alt="figure 3" loading="lazy" width="685" height="561"></picture></a></div><div class="c-article-section__figure-description" data-test="bottom-caption" id="figure-3-desc"><p><b>a</b>, A summary of the number of <i>cupin</i> genes from nine species based on a Pfam search of cupin_1 domain (PF00190). <b>b</b>,<b>c</b>, Phylogenetic trees show <i>cupin</i> genes in nine plant genomes: <i>bicupins</i> (<b>b</b>) and <i>monocupins</i> (<b>c</b>). The colour of each branch corresponds to the background colour for each species in <b>a</b>. The tandem duplicated gene clusters are ordered and shown on scaffolds of the <i>A. angustus</i> genome. The scale bars in the trees show the number of amino acid substitutions per site.</p></div></div><div class="u-text-right u-hide-print"><a class="c-article__pill-button" data-test="article-link" data-track="click" data-track-label="button" data-track-action="view figure" href="/articles/s41477-019-0588-4/figures/3" data-track-dest="link:Figure3 Full size image" aria-label="Full size image figure 3" rel="nofollow"><span>Full size image</span><svg width="16" height="16" focusable="false" role="img" aria-hidden="true" class="u-icon"><use xmlns:xlink="http://www.w3.org/1999/xlink" xlink:href="#icon-eds-i-chevron-right-small"></use></svg></a></div></figure></div></div></div></section><section data-title="CO2-concentrating mechanism"><div class="c-article-section" id="Sec6-section"><h2 class="c-article-section__title js-section-title js-c-reading-companion-sections-item" id="Sec6">CO<sub>2</sub>-concentrating mechanism</h2><div class="c-article-section__content" id="Sec6-content"><p>Hornworts are the only extant land plant lineage harbouring a pyrenoid-based CO<sub>2</sub>-concentrating mechanism (CCM) similar to that of green algae<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 9" title="Villarreal, J. C. & Renzaglia, K. S. The hornworts: important advancements in early land plant evolution. J. Bryol. 37, 157–170 (2015)." href="/articles/s41477-019-0588-4#ref-CR9" id="ref-link-section-d326961673e2470">9</a>,<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 48" title="Hanson, D. T., Renzaglia, K. & Villareal, J. C. in Photosynthesis of Bryophytes and Early Land Plants (eds Hanson, D. T. & Rice, S. K.) 95–111 (Springer, 2014)." href="/articles/s41477-019-0588-4#ref-CR48" id="ref-link-section-d326961673e2473">48</a></sup> (Supplementary Note <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">6.1</a>), for which the key components have been identified<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 49" title="Meyer, M. & Griffiths, H. Origins and diversity of eukaryotic CO2-concentrating mechanisms: lessons for the future. J. Exp. Bot. 64, 769–786 (2013)." href="/articles/s41477-019-0588-4#ref-CR49" id="ref-link-section-d326961673e2480">49</a></sup>. To clarify whether the CCM components of green algae have orthologues in hornworts and other land plants, we searched the <i>A. angustus</i> genome and other plant genomes or transcriptomes with reference to the CCM genes from chlorophyte green algae <i>Chlamydomonas reinhardtii</i><sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 49" title="Meyer, M. & Griffiths, H. Origins and diversity of eukaryotic CO2-concentrating mechanisms: lessons for the future. J. Exp. Bot. 64, 769–786 (2013)." href="/articles/s41477-019-0588-4#ref-CR49" id="ref-link-section-d326961673e2490">49</a>,<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 50" title="Mackinder, L. C. M. A spatial interactome reveals the protein organization of the algal CO2-concentrating mechanism. Cell 171, 133–147 (2017)." href="/articles/s41477-019-0588-4#ref-CR50" id="ref-link-section-d326961673e2493">50</a></sup> (Supplementary Figs. <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">62</a>–<a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">71</a> and Supplementary Note <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">6.2</a>). <i>A. angustus</i> and all other green plants harbour orthologues of <i>CAH1/2</i> whose expression is modulated by external inorganic carbon concentration; of <i>CemA,</i> which maintains stromal pH balance; of <i>LCI11,</i> which mediates the entry of HCO<sup>3−</sup> in the thylakoid lumen; and of <i>RCA1</i> and <i>RBCS1/2</i>, which regulate CO<sub>2</sub> fixation by Rubisco (Supplementary Figs. <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">62</a>, <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">65</a> and <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">69</a>–<a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">72</a>). By contrast, orthologues of <i>CCP1/2</i>, which mediate the entry of HCO<sup>3-</sup> into the chloroplast stroma and of <i>EPYC1</i>, which regulate CO<sub>2</sub> fixation by Rubisco were only present in chlorophyte green algae (Supplementary Figs. <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">67</a> and <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">72</a> and Supplementary Note <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">6.2</a>). The three inorganic carbon transporters (<i>HLA3</i>, <i>LCI1</i> and <i>LCIA</i>-like genes) only occur in bryophytes and green algae, whereas the <i>A. angustus</i> genome lacks the related orthologues (Supplementary Figs. <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">63</a>, <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">66</a> and <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">72</a> and Supplementary Note <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">6.2</a>). Unexpectedly, the three kinds of carbonic anhydrases (CAH3, CAH9 and LCIB/C), which are essential components of CCM, are conserved in non-angiosperm land plants and green algae (Supplementary Figs. <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">62</a>, <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">64</a>, <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">68</a> and <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">72</a>). The <i>A. angustus</i> genome retains the orthologues of both <i>LCIB/C</i> and <i>CAH3</i> genes, but has no copy of <i>CAH9</i> (Supplementary Fig. <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">72</a>). Besides green algae, the essential CCM components occur in both hornworts and other non-angiosperm land plants that lack pyrenoids (Supplementary Fig. <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">72</a>). It implies that the CCM could be an ancestral mechanism of CO<sub>2</sub> fixation by plants, and pyrenoids for CCM are homologous between hornworts and green algae, whereas both CCM components and pyrenoids have undergone multiple losses in land plants in response to atmospheric changes in terrestrial environments<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 10" title="Villarreal, J. C. & Renner, S. S. Hornwort pyrenoids, carbon-concentrating structures, evolved and were lost at least five times during the last 100 million years. Proc. Natl Acad. Sci. USA 109, 18873–18878 (2012)." href="/articles/s41477-019-0588-4#ref-CR10" id="ref-link-section-d326961673e2622">10</a>,<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 48" title="Hanson, D. T., Renzaglia, K. & Villareal, J. C. in Photosynthesis of Bryophytes and Early Land Plants (eds Hanson, D. T. & Rice, S. K.) 95–111 (Springer, 2014)." href="/articles/s41477-019-0588-4#ref-CR48" id="ref-link-section-d326961673e2625">48</a></sup>.</p></div></div></section><section data-title="Horizontal gene transfer"><div class="c-article-section" id="Sec7-section"><h2 class="c-article-section__title js-section-title js-c-reading-companion-sections-item" id="Sec7">Horizontal gene transfer</h2><div class="c-article-section__content" id="Sec7-content"><p>Horizontal gene transfer (HGT) from bacteria or fungi has been reported for both the moss <i>P. patens</i><sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 51" title="Yue, J., Hu, X., Sun, H., Yang, Y. & Huang, J. Widespread impact of horizontal gene transfer on plant colonization of land. Nat. Commun. 3, 1152 (2012)." href="/articles/s41477-019-0588-4#ref-CR51" id="ref-link-section-d326961673e2639">51</a></sup> and the liverwort <i>M. polymorpha</i><sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 15" title="Bowman, J. L. et al. Insights into land plant evolution garnered from the Marchantia polymorpha genome. Cell 171, 287–304 (2017)." href="/articles/s41477-019-0588-4#ref-CR15" id="ref-link-section-d326961673e2645">15</a></sup>. Consistent with those observations, the taxonomic distribution of BLASTP hits following careful phylogenetic analysis and manual inspection suggested that 19 genes from 14 families originated from HGTs from either bacteria or fungi (Supplementary Fig. <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">6</a> and Supplementary Note <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">7.1</a>). Bacterial donors are distributed among nine families: Actinobacteria (three gene families), Alphaproteobacteria (two gene families), Bacteroidetes (two gene families), Firmicutes (one gene family) and Verrucomicrobia (one gene family). Five families were acquired from fungi, belonging to Ascomycota, Basidiomycota, hornwort-symbiotic Chytridiomycota or Mucoromycota<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 13" title="Desirὸ, A., Duckett, J. G., Pressel, S., Villarreal, J. C. & Bidartondo, M. I. Fungal symbioses in hornworts: a chequered history. Proc. R. Soc. B 280, 1759 (2013)." href="/articles/s41477-019-0588-4#ref-CR13" id="ref-link-section-d326961673e2656">13</a></sup> (Fig. <a data-track="click" data-track-label="link" data-track-action="figure anchor" href="/articles/s41477-019-0588-4#Fig4">4a,b</a>, Supplementary Figs. <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">73</a>–<a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">84</a> and <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">86</a> and Supplementary Table <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM3">31</a>). The detection of specific HGT in all three fully sequenced bryophytes is remarkable, and is probably related to the fact that these organisms form symbioses with diverse bacteria and fungi, which, together with the weakly protected tissues in the early developmental stages in the life cycle of these plants, provide the possibility for HGT<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 51" title="Yue, J., Hu, X., Sun, H., Yang, Y. & Huang, J. Widespread impact of horizontal gene transfer on plant colonization of land. Nat. Commun. 3, 1152 (2012)." href="/articles/s41477-019-0588-4#ref-CR51" id="ref-link-section-d326961673e2676">51</a></sup>. In addition, we found that two families originating from HGT from bacteria are shared by the three bryophyte lineages, and one originating from a HGT from fungi is shared between hornworts and liverworts only (Fig. <a data-track="click" data-track-label="link" data-track-action="figure anchor" href="/articles/s41477-019-0588-4#Fig4">4c,d</a>, Supplementary Figs. <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">85</a> and <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">86</a>, Supplementary Table <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM3">31</a> and Supplementary Note <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">7.2</a>). The HGT genes mentioned above (SCUO value 0.2127) exhibit a significantly more biased codon-usage pattern than non-HGT genes (SCUO value 0.1595) (Supplementary Fig. <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">87a</a>), which may be linked to their higher GC content (57.58%) than non-HGT genes (53.26%) (Supplementary Fig. <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">87b</a>).</p><div class="c-article-section__figure js-c-reading-companion-figures-item" data-test="figure" data-container-section="figure" id="figure-4" data-title="Phylogenetic affinities of genes horizontally transferred to A. angustus."><figure><figcaption><b id="Fig4" class="c-article-section__figure-caption" data-test="figure-caption-text">Fig. 4: Phylogenetic affinities of genes horizontally transferred to <i>A. angustus</i>.</b></figcaption><div class="c-article-section__figure-content"><div class="c-article-section__figure-item"><a class="c-article-section__figure-link" data-test="img-link" data-track="click" data-track-label="image" data-track-action="view figure" href="/articles/s41477-019-0588-4/figures/4" rel="nofollow"><picture><source type="image/webp" srcset="//media.springernature.com/lw685/springer-static/image/art%3A10.1038%2Fs41477-019-0588-4/MediaObjects/41477_2019_588_Fig4_HTML.png?as=webp"><img aria-describedby="Fig4" src="//media.springernature.com/lw685/springer-static/image/art%3A10.1038%2Fs41477-019-0588-4/MediaObjects/41477_2019_588_Fig4_HTML.png" alt="figure 4" loading="lazy" width="685" height="615"></picture></a></div><div class="c-article-section__figure-description" data-test="bottom-caption" id="figure-4-desc"><p><b>a</b>, Phylogenetic tree of glyoxalase (PF13468). <b>b</b>, Phylogenetic tree of NAD-binding dehydrogenase (PF08635). <b>c</b>, Phylogenetic tree of glucuronyl hydrolase (PF07470). <b>d</b>, Phylogenetic tree of DNA methyltransferase (PF02870 and PF01035). The stars indicate that the <i>Anthoceros</i> sequence or bryophyte sequences formed a monophyletic clade with homologues of putative HGT donor, reflecting <i>Anthoceros</i>-specific or bryophyte-specific HGT events. Maximum-likelihood bootstrap support values ≥50% are shown above the branches. Red, hornworts and other bryophytes; cyan, green algae; grey, metazoan; orange, stramenopiles; blue, bacteria; yellow, fungi; purple, archaea. The homologues from the kingdom other than the one that HGT donors are involved in are used as the outgroup. The scale bars in the trees show the number of amino acid substitutions per site.</p></div></div><div class="u-text-right u-hide-print"><a class="c-article__pill-button" data-test="article-link" data-track="click" data-track-label="button" data-track-action="view figure" href="/articles/s41477-019-0588-4/figures/4" data-track-dest="link:Figure4 Full size image" aria-label="Full size image figure 4" rel="nofollow"><span>Full size image</span><svg width="16" height="16" focusable="false" role="img" aria-hidden="true" class="u-icon"><use xmlns:xlink="http://www.w3.org/1999/xlink" xlink:href="#icon-eds-i-chevron-right-small"></use></svg></a></div></figure></div><p>The HGT-derived genes in <i>A. angustus</i> mainly contribute to metabolic processes, oxidation–reduction and stress response (Supplementary Table <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM3">31</a>). Some transferred genes related to carbohydrate metabolism are predicted to encode glucuronyl (AANG011893) and glycosyl hydrolases (AANG004297) (Fig. <a data-track="click" data-track-label="link" data-track-action="figure anchor" href="/articles/s41477-019-0588-4#Fig4">4c</a>, Supplementary Fig. <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">79</a> and Supplementary Table <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM3">31</a>), which function in cell wall synthesis and modification and might extend the metabolic flexibility of <i>A. angustus</i> in changing environments<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 52" title="Foflonker, F. et al. Genome of the halotolerant green alga Picochlorum sp. reveals strategies for thriving under fluctuating environmental conditions. Environ. Microbiol. 17, 412–426 (2015)." href="/articles/s41477-019-0588-4#ref-CR52" id="ref-link-section-d326961673e2765">52</a></sup>. The Alphaproteobacteria-derived gene AANG004679 encodes glyoxalase, which is related to drought stress tolerance<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 53" title="Hasanuzzaman, M. et al. Coordinated actions of glyoxalase and antioxidant defense systems in conferring abiotic stress tolerance in plants. Int. J. Mol. Sci. 18, 200 (2017)." href="/articles/s41477-019-0588-4#ref-CR53" id="ref-link-section-d326961673e2769">53</a></sup> (Fig. <a data-track="click" data-track-label="link" data-track-action="figure anchor" href="/articles/s41477-019-0588-4#Fig4">4a</a>). The Actinobacteria-derived DNA methyltransferase genes that are present only in the three groups of bryophytes are related to DNA repair<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 54" title="Finnegan, E. J. & Kovac, K. A. Plant DNA methyltransferases. Plant Mol. Biol. 43, 189–201 (2000)." href="/articles/s41477-019-0588-4#ref-CR54" id="ref-link-section-d326961673e2776">54</a></sup> (Fig. <a data-track="click" data-track-label="link" data-track-action="figure anchor" href="/articles/s41477-019-0588-4#Fig4">4d</a>). The hornworts and liverworts share the fungi-derived terpene synthase-like (<i>MTPSL</i>) genes (Supplementary Fig. <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">85</a>). Terpene synthases are pivotal enzymes for the biosynthesis of terpenoids, which serve as chemical defences against herbivores and pathogens<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 55" title="Jia, Q. et al. Microbial-type terpene synthase genes occur widely in nonseed land plants, but not in seed plants. Proc. Natl Acad. Sci. USA 113, 12328–12333 (2016)." href="/articles/s41477-019-0588-4#ref-CR55" id="ref-link-section-d326961673e2790">55</a></sup>. Some horizontally transferred genes in <i>A. angustus</i>, such as NAD-binding dehydrogenase (Fig. <a data-track="click" data-track-label="link" data-track-action="figure anchor" href="/articles/s41477-019-0588-4#Fig4">4b</a>) and <i>MTPSL</i> genes (Supplementary Fig. <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">85</a>), underwent subsequent gene duplications. The results suggest that the acquisition of foreign genes might have provided additional means for environmental adaptation during evolution of the hornwort lineage.</p></div></div></section><section data-title="Conclusions"><div class="c-article-section" id="Sec8-section"><h2 class="c-article-section__title js-section-title js-c-reading-companion-sections-item" id="Sec8">Conclusions</h2><div class="c-article-section__content" id="Sec8-content"><p>As land pioneers, the three bryophyte groups form a well-supported monophyletic lineage, with hornworts sister to liverworts and mosses. The genome of hornwort <i>A. angustus</i> shows no evidence of WGDs and low genetic redundancy for networks underlying plant body plan, which may be congruent with an overall simple body plan. Hornworts have retained the essential components of CCM found in green algae in response to the atmospheric changes in terrestrial environments. Meanwhile, the gene inventory in <i>A. angustus</i> expanded mainly through tandem duplication and HGT. In particular, the expansion of specific gene families and the acquisition of foreign genes have provided additional metabolic abilities in hornworts that probably facilitated their survival in a terrestrial environment. Together, our results indicate how the draft genome of <i>A. angustus</i> provides a useful model for studying early land plant evolution and the mechanism of plant terrestrial adaptation.</p></div></div></section><section data-title="Methods"><div class="c-article-section" id="Sec9-section"><h2 class="c-article-section__title js-section-title js-c-reading-companion-sections-item" id="Sec9">Methods</h2><div class="c-article-section__content" id="Sec9-content"><h3 class="c-article__sub-heading" id="Sec10">Sample preparation and sequencing</h3><p>The natural populations of <i>A. angustus</i> Steph. were collected from Jinping County, Yunnan Province, China. The voucher specimen has been deposited at the herbarium, Institute of Botany, Chinese Academy of Sciences, Beijing, China with collection number W1879-2010-01-18. The sporophytes of <i>A. angustus</i> were detached from the gametophytes, sterilized in 10% sodium hypochlorite and subsequently rinsed with distilled water<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 56" title="Duckett, J. G. et al. In vitro cultivation of bryophytes: a review of practicalities, problems, progress and promise. J. Bryol. 26, 3–20 (2004)." href="/articles/s41477-019-0588-4#ref-CR56" id="ref-link-section-d326961673e2842">56</a></sup>. The sporangium was opened and the spores were homogenized and spread onto the 1/2 KnopII agar medium<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 57" title="Kugita, M. et al. The complete nucleotide sequence of the hornwort (Anthoceros formosae) chloroplast genome: insight into the earliest land plants. Nucleic Acids Res. 31, 716–721 (2003)." href="/articles/s41477-019-0588-4#ref-CR57" id="ref-link-section-d326961673e2846">57</a></sup> in Petri dishes (Supplementary Fig. <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">1b</a>). The culture temperature was between 21 °C and 25 °C. Spores germinated within a couple of days, and then the sporelings started to grow. After approximately three to four weeks, the gametophyte started to grow (Supplementary Fig. <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">1c,d</a>). Since spores are aposymbiotic, we did not find the phenomenon of mucilage-filled cavities colonization by cyanobacteria on the <i>A. angustus</i> gametophyte during the sterile culture. A gametophyte from a single spore was selected and cultured by asexual propagation. The tissue yielded from subculture was used for genome and RNA sequencing. We tried to induce sexual reproduction by dropping the growth temperature of gametophyte cultures to 10 °C and 16 °C, respectively; however until now they have not yet produced reproductive organs. Therefore, the sequenced <i>A. angustus</i> is indeed a single-sex individual, which is sequenced at the gametophyte phase of its life cycle.</p><p>Genomic DNA was isolated using the Plant DNAzol reagent for genomic DNA extraction (Life Technologies) according to the manufacturer’s protocols. For whole-genome shotgun sequencing, ten sequencing libraries with insert sizes ranging from 170 bp to 40 kb were generated (Supplementary Table <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">1</a>). Sequencing libraries were constructed using a library construction kit (Illumina). All libraries were sequenced on the Illumina HiSeq 2000 platform. Raw sequencing reads were trimmed with Trimmomatic (v.0.33)<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 58" title="Bolger, A. M., Lohse, M. & Usadel, B. Trimmomatic: a flexible trimmer for Illumina sequence data. Bioinformatics 30, 2114–2120 (2014)." href="/articles/s41477-019-0588-4#ref-CR58" id="ref-link-section-d326961673e2869">58</a></sup>. Only high-quality reads with a total length of 126,532,381,412 bp were used for further analysis (Supplementary Table <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">1</a>). For Oxford Nanopore sequencing, we constructed a genomic DNA library using the ONT 1D ligation sequencing kit (SQK-LSK108) according to the manufacturer’s instructions. The sequencing used a single 1D flow cell on a PromethION sequencer (Oxford Nanopore Technologies). A total of 63,614,292,295 bp raw reads were generated, of which 36,070,452,175 bp were retained for further analysis after filtering and trimming (Supplementary Table <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">3</a>).</p><p>Total RNA was extracted using the PureLink Plant RNA reagent (Life Technologies) and further purified using TRIzol reagent (Invitrogen). For transcriptome sequencing (RNA sequencing), libraries with insert sizes ranging from 200 bp to 500 bp were constructed using the mRNA-Seq Prep Kit (Illumina) and then sequenced using the Illumina HiSeq 2000 platform. For small-RNA sequencing, the library was generated from RNA sample using the Truseq Small RNA Preparation kit (Illumina) and sequenced on the Illumina HiSeq 2500 platform.</p><h3 class="c-article__sub-heading" id="Sec11">Decontamination</h3><p>The GC content versus <i>k</i>-mer frequency distribution pattern of the Illumina raw reads (Supplementary Table <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">1</a>) after trimming presented two large groups: one group with a low <i>k</i>-mer frequency (<50) and a wide GC content distribution range (median number at 0.7), and the other group with a high <i>k</i>-mer frequency (60–165) and a concentrated GC content distribution range (median number at 0.5) (Supplementary Fig. <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">2a</a>). The BLASTN results against the NCBI nucleotide database revealed that the former sequences were mainly from a variety of bacteria and the latter were the real genome sequences of <i>A. angustus</i>. We also investigated the <i>k</i>-mer distributions of the raw reads from the other two published hornwort genomic sequences, <i>A. agrestis</i> (accession: ERX714368)<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 59" title="Szövényi, P. et al. Establishment of Anthoceros agrestis as a model species for studying the biology of hornworts. BMC Plant Biol. 15, 98 (2015)." href="/articles/s41477-019-0588-4#ref-CR59" id="ref-link-section-d326961673e2915">59</a></sup> and <i>Anthoceros punctatus</i> (accession: <a href="https://www.ncbi.nlm.nih.gov/sra/?term=SRX538621">SRX538621</a>)<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 60" title="Li, F. et al. Horizontal transfer of an adaptive chimeric photoreceptor from bryophytes to ferns. Proc. Natl Acad. Sci. USA 111, 6672–6677 (2014)." href="/articles/s41477-019-0588-4#ref-CR60" id="ref-link-section-d326961673e2930">60</a></sup>, and found a similar distribution pattern as that of <i>A. angustus</i>, containing two groups, one for the contaminant sequences and the other for sequences of the plant itself (Supplementary Fig. <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">2c,d</a>). Because external bacterial contaminations from the laboratory cause <i>A. angustus</i> to turn yellow and die during culturing, and all three <i>Anthoceros</i> species through axenic cultures still have the same bacterial contamination problems (Supplementary Fig. <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">2a,c,d</a>), we infer that these bacterial contaminations are from symbiotic bacteria of <i>Anthoceros</i> that might accompany spores hiding in the sterilized sporangium. Furthermore, we performed the DAPI staining analysis<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 61" title="Mergaert, P. et al. Eukaryotic control on bacterial cell cycle and differentiation in the Rhizobium–legume symbiosis. Proc. Natl Acad. Sci. USA 103, 5230–5235 (2006)." href="/articles/s41477-019-0588-4#ref-CR61" id="ref-link-section-d326961673e2953">61</a></sup> to investigate the distribution of symbiotic bacteria in <i>A. angustus</i>. The gametophytes were stained by 0.2 mg l<sup>−1</sup> DAPI (4′,6-diamidino-2-phenylindole dihydrochloride; Sigma, cat. no. D9564) for five minutes. The stained gametophytes were washed three times, and then observed using confocal microscopy. The bacterial micro-colonies were observed on the outer surface, as well as in the intercellular space of the gametophytes of <i>A. angustus</i> (Supplementary Fig. <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">3</a>). Based on the GC content versus <i>k</i>-mer frequency distribution pattern of the Illumina raw reads and the result of the DAPI staining, we could imagine that there is a certain amount of bacterial sequences remaining in the genome sequencing data of <i>A. angustus</i>. In order to isolate them, we performed a series of decontamination steps. After generating the <i>k</i>-mer frequency, we chose the high-abundance <i>k</i>-mer depth (60–165) and retained the corresponding reads for further analysis. This treatment yielded filtered reads with a total length of 17,099,027,576 bp (Supplementary Table <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">2</a>). The distribution pattern of GC content versus <i>k</i>-mer frequency of the <i>A. angustus</i> filtered reads is depicted in Supplementary Fig. <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">2b</a>, which shows an entire group with a sequencing depth of approximately 150×. Furthermore, we performed error correction for filtered Nanopore reads using decontaminated Illumina reads by Nextdenovo (v.2.0)<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 62" title="Koren, S. et al. Canu: scalable and accurate long-read assembly via adaptive k-mer weighting and repeat separation. Genome Res. 27, 722–736 (2017)." href="/articles/s41477-019-0588-4#ref-CR62" id="ref-link-section-d326961673e2994">62</a></sup>, resulting in 9,247,957,448 bp corrected reads (Supplementary Table <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">3</a>). Through MEGABLAST against the NCBI nucleotide database, we further removed 5,463,972,682 bp prokaryotic sequences or organellar sequences, and finally got 3,783,984,766 clean reads with a sequencing depth of approximately 35× (Supplementary Table <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">3</a>). A total of approximately 185× coverage was obtained finally.</p><h3 class="c-article__sub-heading" id="Sec12">Genome size estimation</h3><p>To estimate the genome size of <i>A. angustus</i>, we used clean Illumina reads to calculate the <i>k</i>-mer distribution. According to the Lander–Waterman theory<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 63" title="Arratia, R., Lander, E. S., Tavaré, S. & Waterman, M. S. Genomic mapping by anchoring random clones: a mathematical analysis. Genomics 11, 806–827 (1991)." href="/articles/s41477-019-0588-4#ref-CR63" id="ref-link-section-d326961673e3018">63</a></sup>, the genome size can be determined by dividing the total number of <i>k</i>-mers by the peak value of the <i>k</i>-mer distribution. Because we sequenced the haploid gametophyte of <i>A. angustus</i>, only one peak was found in the <i>k</i>-mer distribution. The total number of <i>k</i>-mers was 14,092,039,150, and the position of the peak was at 132 (Supplementary Fig. <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">4</a>). The peak was used as the expected <i>k</i>-mer depth and substituted into the formula genome size = total <i>k</i>-mer/expected <i>k</i>-mer depth, and the haploid genome size was estimated to be 106,757,872 bp (Supplementary Fig. <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">4</a>).</p><h3 class="c-article__sub-heading" id="Sec13">Genome assembly and assessment</h3><p>The clean Nanopore reads after filtering and decontamination were assembled with wtdbg-1.2.8. After finishing the pre-assembly (148 Mb), iterative polishing was conducted using Pilon (v.1.22)<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 64" title="Walker, B. J. et al. Pilon: an integrated tool for comprehensive microbial variant detection and genome assembly improvement. PLoS ONE 9, e112963 (2014)." href="/articles/s41477-019-0588-4#ref-CR64" id="ref-link-section-d326961673e3062">64</a></sup> in which clean Illumina reads were aligned with the pre-assembled contigs. The pre-assembled contig sequences were performed with the MEGABLAST search against the NCBI nucleotide database to further remove prokaryotic sequences or organellar DNA. A total of approximately 29 Mb of data were removed. Further, we combined the final pre-assembled contig sequences from Nanopore sequencing and clean paired-read data from Illumina sequencing into scaffolds using SSPACE (v.3.0)<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 65" title="Boetzer, M., Henkel, C. V., Jansen, H. J., Butler, D. & Pirovano, W. Scaffolding pre-assembled contigs using SSPACE. Bioinformatics 27, 578–579 (2011)." href="/articles/s41477-019-0588-4#ref-CR65" id="ref-link-section-d326961673e3066">65</a></sup> tool (Supplementary Table <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">4</a>). Genome assembly completeness was assessed using the plantae database of 956 single-copy orthologues using BUSCO (v.3)<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 16" title="Simao, F. A., Waterhouse, R. M., Ioannidis, P., Kriventseva, E. V. & Zdobnov, E. M. BUSCO: assessing genome assembly and annotation completeness with single-copy orthologs. Bioinformatics 31, 3210–3212 (2015)." href="/articles/s41477-019-0588-4#ref-CR16" id="ref-link-section-d326961673e3073">16</a></sup> with a BLAST threshold <i>E</i>-value of 1 × 10<sup>−5</sup> (Supplementary Table <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">10</a>).</p><h3 class="c-article__sub-heading" id="Sec14">Transcriptome assembly and mapping</h3><p>We used Trimmomatic<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 58" title="Bolger, A. M., Lohse, M. & Usadel, B. Trimmomatic: a flexible trimmer for Illumina sequence data. Bioinformatics 30, 2114–2120 (2014)." href="/articles/s41477-019-0588-4#ref-CR58" id="ref-link-section-d326961673e3095">58</a></sup> to remove adaptors from the raw reads of transcriptome sequences and filter out low-quality reads before assembly. The resulting high-quality reads were de novo assembled and annotated using Trinity (v.2.5.1)<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 66" title="Haas, B. J. et al. De novo transcript sequence reconstruction from RNA-seq using the Trinity platform for reference generation and analysis. Nat. Protoc. 8, 1494–1512 (2013)." href="/articles/s41477-019-0588-4#ref-CR66" id="ref-link-section-d326961673e3099">66</a></sup>. For genes with more than one transcript, the longest transcript was chosen as the unigene and used to predict open reading frames (ORFs) using TransDecoder (v.5.0.2) (<a href="https://github.com/TransDecoder/TransDecoder/wiki">https://github.com/TransDecoder/TransDecoder/wiki</a>). Finally, we obtained 39,044 unigenes, 26,805 of which had predicted ORFs. To extend the validation of genome assembly, the transcriptome was compared to the reference assembly using BLASTN, with an <i>E</i>-value <1 × 10<sup>−5</sup>. Of the 26,805 transcripts (>200 bp), 97.66% were successfully mapped back to the final assembled genome (Supplementary Table <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">5</a>).</p><h3 class="c-article__sub-heading" id="Sec15">Repeat prediction</h3><p>Tandem Repeats Finder (v.4.09)<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 67" title="Benson, G. Tandem repeats finder: a program to analyze DNA sequences. Nucleic Acids Res. 27, 573–580 (1999)." href="/articles/s41477-019-0588-4#ref-CR67" id="ref-link-section-d326961673e3127">67</a></sup> was used to search for tandem repeats in the <i>A. angustus</i> genome. Both homology-based and de novo approaches were used to search for TEs. In the homology-based approach, we used RepeatMasker (v.4.1.0)<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 67" title="Benson, G. Tandem repeats finder: a program to analyze DNA sequences. Nucleic Acids Res. 27, 573–580 (1999)." href="/articles/s41477-019-0588-4#ref-CR67" id="ref-link-section-d326961673e3134">67</a></sup> and RepeatProteinMask<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 68" title="Tarailo-Graovac, M. & Chen, N. Using RepeatMasker to identify repetitive elements in genomic sequences. Curr. Protoc. Bioinformatics 25, 4.10.1–4.10.14 (2009)." href="/articles/s41477-019-0588-4#ref-CR68" id="ref-link-section-d326961673e3138">68</a></sup> with the Repbase<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 69" title="Jurka, J. et al. Repbase update, a database of eukaryotic repetitive elements. Cytogenet. Genome Res. 110, 462–467 (2005)." href="/articles/s41477-019-0588-4#ref-CR69" id="ref-link-section-d326961673e3142">69</a></sup> database of known repeat sequences to search for the TEs in the <i>A. angustus</i> genome. In the de novo approach, we used LTR_FINDER (v.1.0.2)<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 70" title="Xu, Z. & Wang, H. LTR_FINDER: an efficient tool for the prediction of full-length LTR retrotransposons. Nucleic Acids Res. 35, W265–W268 (2007)." href="/articles/s41477-019-0588-4#ref-CR70" id="ref-link-section-d326961673e3150">70</a></sup>, PILER (v.1.3.4.)<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 71" title="Edgar, R. C. & Myers, E. W. PILER: identification and classification of genomic repeats. Bioinformatics 21(Suppl. 1), i152–i158 (2005)." href="/articles/s41477-019-0588-4#ref-CR71" id="ref-link-section-d326961673e3154">71</a></sup> and RepeatModeler (v.1.0.3)<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 72" title="Price, A. L., Jones, N. C. & Pevzner, P. A. De novo identification of repeat families in large genomes. Bioinformatics 21(Suppl. 1), i351–i358 (2005)." href="/articles/s41477-019-0588-4#ref-CR72" id="ref-link-section-d326961673e3158">72</a></sup> to construct a de novo repeat sequence database for <i>A. angustus</i> and then used RepeatMasker to search for repeats in the genome. All the repeats identified by different methods were combined into the final repeat annotation after removing the redundant repeats. The predicted repeats covered 64.21% of the genome sequence (Supplementary Table <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">6</a>). The categories of predicted TEs in the <i>A. angustus</i> genome are summarized in Supplementary Table <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">7</a>.</p><h3 class="c-article__sub-heading" id="Sec16">Genome annotation</h3><p>To predict protein-coding genes, three approaches were used: (1) de novo gene prediction, (2) homology-based prediction, and (3) RNA-sequencing annotation. For de novo prediction, AUGUSTUS (v.2.5.5)<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 73" title="Stanke, M. et al. AUGUSTUS: ab initio prediction of alternative transcripts. Nucleic Acids Res. 34, W435–W439 (2006)." href="/articles/s41477-019-0588-4#ref-CR73" id="ref-link-section-d326961673e3183">73</a></sup> and GlimmerHMM (v3.0.1)<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 74" title="Majoros, W. H., Pertea, M. & Salzberg, S. L. TigrScan and GlimmerHMM: two open source ab initio eukaryotic gene-finders. Bioinformatics 20, 2878–2879 (2004)." href="/articles/s41477-019-0588-4#ref-CR74" id="ref-link-section-d326961673e3187">74</a></sup> were applied to predict genes. For homology-based prediction, we mapped the protein sequences of five published green plant genomes (<i>Arabidopsis thaliana</i>, <i>Selaginella moellendorffii</i>, <i>P. patens</i>, <i>M. polymorpha</i> and <i>Klebsormidium nitens</i>) onto the <i>A. angustus</i> genome using TBLASTN, with a threshold <i>E</i>-value of 1 × 10<sup>−5</sup>, and then used GeneWise (v.2.4.1)<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 75" title="Birney, E., Clamp, M. & Durbin, R. GeneWise and Genomewise. Genome Res. 14, 988–995 (2004)." href="/articles/s41477-019-0588-4#ref-CR75" id="ref-link-section-d326961673e3215">75</a></sup> to predict gene structures. The de novo set and five homologue-based results were combined by MAKER (v.1.0)<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 76" title="Holt, C. & Yandell, M. MAKER2: an annotation pipeline and genome-database management tool for second-generation genome projects. BMC Bioinf. 12, 491 (2011)." href="/articles/s41477-019-0588-4#ref-CR76" id="ref-link-section-d326961673e3220">76</a></sup> to integrate a consensus gene set (Supplementary Table <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">8</a>). To supplement and improve the gene set, we aligned the RNA-sequencing data to the genome using TopHat (v2.1.1)<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 77" title="Trapnell, C., Pachter, L. & Salzberg, S. L. TopHat: discovering splice junctions with RNA-seq. Bioinformatics 25, 1105–1111 (2009)." href="/articles/s41477-019-0588-4#ref-CR77" id="ref-link-section-d326961673e3227">77</a></sup>, and the alignments were used as input for Cufflinks (v.2.2.1)<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 78" title="Trapnell, C. et al. Differential gene and transcript expression analysis of RNA-seq experiments with TopHat and Cufflinks. Nat. Protoc. 7, 562–578 (2012)." href="/articles/s41477-019-0588-4#ref-CR78" id="ref-link-section-d326961673e3231">78</a></sup> with default parameters. We manually combined the MAKER gene set and ORFs of transcripts to form the final gene set that contains 14,629 genes (Supplementary Table <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">8</a>).</p><p>The <i>A. angustus</i> predicted genes were aligned against the sequences in NCBI non-redundant protein database using BLASTP<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 79" title="Camacho, C. et al. BLAST+: architecture and applications. BMC Bioinf. 10, 421 (2009)." href="/articles/s41477-019-0588-4#ref-CR79" id="ref-link-section-d326961673e3244">79</a></sup> (<i>E</i>-value <1 × 10<sup>−5</sup>). According to the NCBI taxonomy categories of best BLAST hits, the source of <i>A. angustus</i> genes were classified (Supplementary Fig. <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">6</a>). Functional annotation of these predicted genes was obtained by aligning the protein sequences of these genes against the sequences in public protein databases using BLASTP<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 79" title="Camacho, C. et al. BLAST+: architecture and applications. BMC Bioinf. 10, 421 (2009)." href="/articles/s41477-019-0588-4#ref-CR79" id="ref-link-section-d326961673e3260">79</a></sup> (<i>E</i>-value <1 × 10<sup>−5</sup>, identity >30% and coverage >70%, excluding annotations only characterized as hypothetical or predicted protein), including, SwissProt<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 80" title="Boeckmann, B. et al. The SWISS-PROT protein knowledgebase and its supplement TrEMBL in 2003. Nucleic Acids Res. 31, 365–370 (2003)." href="/articles/s41477-019-0588-4#ref-CR80" id="ref-link-section-d326961673e3269">80</a></sup>, TrEMBL<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 80" title="Boeckmann, B. et al. The SWISS-PROT protein knowledgebase and its supplement TrEMBL in 2003. Nucleic Acids Res. 31, 365–370 (2003)." href="/articles/s41477-019-0588-4#ref-CR80" id="ref-link-section-d326961673e3273">80</a></sup>, Pfam<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 81" title="Finn, R. D. et al. Pfam: the protein families database. Nucleic Acids Res. 42, D222–D230 (2014)." href="/articles/s41477-019-0588-4#ref-CR81" id="ref-link-section-d326961673e3278">81</a></sup>, GO<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 82" title="Ashburner, M. et al. Gene ontology: tool for the unification of biology. Nat. Genet. 25, 25–29 (2000)." href="/articles/s41477-019-0588-4#ref-CR82" id="ref-link-section-d326961673e3282">82</a></sup> and KEGG<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 83" title="Kanehisa, M. & Goto, S. KEGG: Kyoto encyclopedia of genes and genomes. Nucleic Acids Res. 28, 27–30 (2000)." href="/articles/s41477-019-0588-4#ref-CR83" id="ref-link-section-d326961673e3286">83</a></sup> (Supplementary Tables <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">9</a> and <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM3">32</a>).</p><h3 class="c-article__sub-heading" id="Sec17">Identification of non-coding RNA genes</h3><p>To obtain a reliable profile of <i>A. angustus</i> miRNAs, we used mapped reads from small-RNA sequencing with reference to the <i>A. angustus</i> draft genome to search against miRNA sequences in <i>A. thaliana</i>, <i>Oryza sativa</i>, <i>S. moellendorffii</i>, <i>P. patens</i> and <i>C. reinhardtii</i> from miRBase (<a href="http://www.mirbase.org/">http://www.mirbase.org/</a>) for predicting the known miRNAs. The mapped reads were also used to identify novel miRNAs using miREvo (v.1.2)<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 84" title="Wen, M., Shen, Y., Shi, S. & Tang, T. miREvo: an integrative microRNA evolutionary analysis platform for next-generation sequencing experiments. BMC Bioinf. 13, 140 (2012)." href="/articles/s41477-019-0588-4#ref-CR84" id="ref-link-section-d326961673e3333">84</a></sup> software. The tRNA genes were searched by tRNAscan-SE (v.1.3.1)<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 85" title="Lowe, T. M. & Eddy, S. R. tRNAscan-SE: a program for improved detection of transfer RNA genes in genomic sequence. Nucleic Acids Res. 25, 955–964 (1997)." href="/articles/s41477-019-0588-4#ref-CR85" id="ref-link-section-d326961673e3337">85</a></sup>. The rRNA genes were predicted by aligning plant rRNA sequences from NCBI (<i>A. thaliana</i> and <i>Anthoceros agrestis</i>) to the <i>A. angustus</i> genome by BLASTN. The snRNA genes were predicted using INFERNAL (v.1.1)<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 86" title="Nawrocki, E. P. & Eddy, S. R. Infernal 1.1: 100-fold faster RNA homology searches. Bioinformatics 29, 2933–2935 (2013)." href="/articles/s41477-019-0588-4#ref-CR86" id="ref-link-section-d326961673e3351">86</a></sup> to search from the Rfam database.</p><h3 class="c-article__sub-heading" id="Sec18">Gene-family identification</h3><p>To construct the dataset for gene-family clustering, the protein-coding genes from the genomes of <i>A. angustus</i> and 18 other green plants were used, including those of seven angiosperms (<i>A. thaliana</i>, <i>Genlisea aurea</i>, <i>Vitis vinifera</i>, <i>O. sativa</i>, <i>Phalaenopsis equestris</i>, <i>Zostera marina</i> and <i>Amborella trichopoda</i>), one gymnosperm (<i>Picea abies</i>), one lycophyte (<i>S. moellendorffii</i>), two bryophytes (moss <i>P. patens</i> and liverwort <i>M. polymorpha</i>), two charophytes (<i>Chara braunii</i> and <i>K. nitens</i>) and five chlorophytes (<i>Volvox carteri</i>, <i>Chlamydomonas reinhardtii</i>, <i>Ulva mutabilis</i>, <i>Coccomyxa subellipsoidea</i> and <i>Chlorella variabilis</i>) (Supplementary Table <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">13</a>). We chose the longest transcript to represent each gene and removed mitochondrial and chloroplast genes. After performing an all-against-all BLASTP search with a threshold <i>E</i>-value of 1 × 10<sup>−5</sup>, identity >30% and coverage >30%, orthogroups or putative gene families or subfamilies were identified using OrthoMCL (v.2.0)<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 87" title="Li, L., Stoeckert, C. J. & Roos, D. S. OrthoMCL: identification of ortholog groups for eukaryotic genomes. Genome Res. 13, 2178–2189 (2003)." href="/articles/s41477-019-0588-4#ref-CR87" id="ref-link-section-d326961673e3431">87</a></sup>, on the basis of a collection of 397,132 predicted protein-coding genes from the above 19 Viridiplantae genomes. A 5-way comparison of <i>A. angustus</i>, Setaphyta (<i>M. polymorpha</i> and <i>P. patens</i>), Tracheophyta (vascular plants) (<i>A. thaliana</i>, <i>V. vinifera</i>, <i>O. sativa</i>, <i>Z. marina</i>, <i>P. equestris</i>, <i>A. trichopoda</i>, <i>P. abies</i>, <i>G. aurea</i> and <i>S. moellendorffii</i>), Charophyta (<i>C. braunii</i> and <i>K. nitens</i>) and Chlorophyta (<i>V. carteri</i>, <i>C. reinhardtii</i>, <i>U. mutabilis</i>, <i>C. subellipsoidea</i> and <i>C. variabilis</i>) is shown in Fig. <a data-track="click" data-track-label="link" data-track-action="figure anchor" href="/articles/s41477-019-0588-4#Fig1">1a</a>. For <i>A. angustus</i>-specific gene families, we conducted GO and KEGG enrichment analyses via an enrichment pipeline (<a href="https://sourceforge.net/projects/enrichmentpipeline/">https://sourceforge.net/projects/enrichmentpipeline/</a>).</p><h3 class="c-article__sub-heading" id="Sec19">Phylogenomics</h3><p>We extracted 85 single-copy gene families shared by 19 Viridiplantae for phylogenomic analysis (Supplementary Note <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">2.1</a>). The amino acid alignments of each single-copy gene family were aligned by MAFFT (v.7)<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 88" title="Katoh, K., Kuma, K. I., Toh, H. & Miyata, T. MAFFT version 5: improvement in accuracy of multiple sequence alignment. Nucleic Acids Res. 33, 511–518 (2005)." href="/articles/s41477-019-0588-4#ref-CR88" id="ref-link-section-d326961673e3520">88</a></sup>, and the nucleotide alignments were generated separately with TranslatorX (v0.9)<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 89" title="Abascal, F., Zardoya, R. & Telford, M. J. TranslatorX: multiple alignment of nucleotide sequences guided by amino acid translations. Nucleic Acids Res. 38, W7–W13 (2010)." href="/articles/s41477-019-0588-4#ref-CR89" id="ref-link-section-d326961673e3524">89</a></sup> on the basis of the corresponding amino acid translation. The amino acid data, the complete nucleotide data and the first and second codon positions, as well as the third codon positions, were concatenated as super-matrices. These data matrices were used for maximum likelihood phylogenetic analyses by RAxML (v.7.2.3)<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 90" title="Stamatakis, A. RAxML-VI-HPC: maximum likelihood-based phylogenetic analyses with thousands of taxa and mixed models. Bioinformatics 22, 2688–2690 (2006)." href="/articles/s41477-019-0588-4#ref-CR90" id="ref-link-section-d326961673e3528">90</a></sup> with the GTR + Γ and JTT models for nucleotide and amino acid data, respectively. For each analysis, the bootstrap support was estimated based on 300 pseudoreplicates using a GTR + CAT approximation. To estimate the degree of substitutional saturation for the four concatenated datasets mentioned above (Supplementary Note <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">2.2</a>), we plotted the uncorrected <i>p</i>-distances against the inferred distances using the method described by Forterre and Philippe<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 91" title="Forterre, P. & Philippe, H. Where is the root or the universal tree of life? Bioessays 21, 871–879 (1999)." href="/articles/s41477-019-0588-4#ref-CR91" id="ref-link-section-d326961673e3539">91</a></sup>. The level of saturation was estimated by computing the slope of the regression line in the plot; the shallower the slope, the greater the degree of saturation. The maximum composite likelihood method was used to calculate the inferred distances for nucleotide data and Poisson correction was used to calculate the inferred distances for the amino acid data.</p><p>To improve the taxon sampling in bryophytes for divergence time estimation, the transcriptome sequences of 22 other bryophytes were downloaded from the 1KP database<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 92" title="Matasci, N. et al. Data access for the 1,000 Plants (1KP) project. Gigascience 3, 17 (2014)." href="/articles/s41477-019-0588-4#ref-CR92" id="ref-link-section-d326961673e3546">92</a></sup> (<a href="http://www.onekp.com/public_data.html">http://www.onekp.com/public_data.html</a>) and used in subsequent analyses (Supplementary Table <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">18</a> and Supplementary Note <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">2.3</a>). The divergence time was estimated using the MCMCTree program in the PAML package (v.4.7)<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 93" title="Yang, Z. PAML 4: phylogenetic analysis by maximum likelihood. Mol. Biol. Evol. 24, 1586–1591 (2007)." href="/articles/s41477-019-0588-4#ref-CR93" id="ref-link-section-d326961673e3563">93</a></sup> under the nucleotide general time reversible (GTR) substitution model and with the independent rate model as the molecular clock model. The Markov chain Monte Carlo (MCMC) process consists of 500,000 burn-in iterations and 1,500,000 sampling iterations (1 sample per 150 iterations). The same parameters were executed twice to obtain a stable result. We applied nine node constraints in the age estimate (Supplementary Fig. <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">10</a>). The minimum and maximum constraints for each node are shown in Supplementary Table <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">19</a>.</p><p>Gene-family sizes were inferred from the gene-family profile obtained by the program OrthoMCL. The minimum ancestral gene families were estimated using DOLLOP program included in the PHYLIP package (v.3.695)<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 94" title="Felsenstein, J. PHYLIP: phylogenetic inference program v.3.6 (Univ. of Washington, 2005)." href="/articles/s41477-019-0588-4#ref-CR94" id="ref-link-section-d326961673e3577">94</a></sup> to determine gene-family gain or loss evolutions of gene families. There are 8,141 gene families in the <i>A. angustus</i> genome, 8,944 in <i>M. polymorpha</i> and 9,566 in <i>P. patens</i>, and 9,789 ancestral families in the ancestral bryophyte lineage (Fig. <a data-track="click" data-track-label="link" data-track-action="figure anchor" href="/articles/s41477-019-0588-4#Fig1">1b</a>).</p><h3 class="c-article__sub-heading" id="Sec20"><i>K</i> <sub>S</sub> distribution and co-linearity analysis</h3><p>All <i>K</i><sub>S</sub> distributions were constructed using wgd (v.3.0)<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 95" title="Zwaenepoel, A. & Van de Peer, Y. wgd—simple command line tools for the analysis of ancient whole genome duplications. Bioinformatics 35, 2153–2155 (2018)." href="/articles/s41477-019-0588-4#ref-CR95" id="ref-link-section-d326961673e3611">95</a></sup> using default settings. The <i>M. polymorpha</i> and <i>P. patens</i> genome data was acquired from the PLAZA resource<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 96" title="Van Bel, M. et al. PLAZA 4.0: an integrative resource for functional, evolutionary and comparative plant genomics. Nucleic Acids Res. 46, D1190–D1196 (2018)." href="/articles/s41477-019-0588-4#ref-CR96" id="ref-link-section-d326961673e3621">96</a></sup>. Pairwise co-linearity analyses within and between <i>A</i>. <i>angustus</i>, <i>M. polymorpha</i> and <i>P. patens</i> were conducted using I-ADHoRe 3.0<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 97" title="Proost, S. et al. i-ADHoRe 3.0―fast and sensitive detection of genomic homology in extremely large data sets. Nucleic Acids Res. 40, e11 (2012)." href="/articles/s41477-019-0588-4#ref-CR97" id="ref-link-section-d326961673e3638">97</a></sup> with the following parameter settings: gap_size = 30, cluster_gap = 35, q_value = 0.75, prob_cutoff = 0.01, anchor_points = 3, alignment_method = ‘gg2’, level_2_only = ‘false’, table_type = ‘family’ and multiple_hypothesis_correction = ‘FDR’. Within-genome co-linearity analyses were based on the paralogous families inferred with wgd, whereas the between-genome co-linearity analyses were conducted using gene families inferred with OrthoFinder using default settings.</p><h3 class="c-article__sub-heading" id="Sec21">Analysis of TFs</h3><p>We used the genome-wide TF prediction program iTAK (v.1.7)<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 98" title="Zheng, Y. et al. iTAK: a program for genome-wide prediction and classification of plant transcription factors, transcriptional regulators, and protein kinases. Mol. Plant 9, 1667–1670 (2016)." href="/articles/s41477-019-0588-4#ref-CR98" id="ref-link-section-d326961673e3650">98</a></sup> (<a href="http://bioinfo.bti.cornell.edu/cgi-bin/itak/index.cgi">http://bioinfo.bti.cornell.edu/cgi-bin/itak/index.cgi</a>) with default parameters to preliminarily identify TFs in the above 19 Viridiplantae (Supplementary Tables <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">13</a> and <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM3">22</a>). The reconstruction of the ancestral state for the individual TF family was performed using Mesquite (v.3.51)<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 99" title="Maddison, W. P. & Maddison, D. R. Mesquite: a modular system for evolutionary analysis v.2.75 (Mesquite Project, 2011)." href="/articles/s41477-019-0588-4#ref-CR99" id="ref-link-section-d326961673e3667">99</a></sup> (<a href="http://mesquiteproject.org/">http://mesquiteproject.org/</a>), and the most parsimonious assumption was taken.</p><h3 class="c-article__sub-heading" id="Sec22">Phylogenetic analysis of gene families</h3><p>Generally, HMMER search<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 100" title="Madera, M. & Gough, J. A comparison of profile hidden Markov model procedures for remote homology detection. Nucleic Acids Res. 30, 4321–4328 (2002)." href="/articles/s41477-019-0588-4#ref-CR100" id="ref-link-section-d326961673e3687">100</a></sup> with a domain profile or BLAST search using known protein sequences from other plants as queries was performed to retrieve the sequences from the <i>A. angustus</i> genome (Supplementary Notes <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">4–6</a>). The results of TF prediction by iTAK<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 98" title="Zheng, Y. et al. iTAK: a program for genome-wide prediction and classification of plant transcription factors, transcriptional regulators, and protein kinases. Mol. Plant 9, 1667–1670 (2016)." href="/articles/s41477-019-0588-4#ref-CR98" id="ref-link-section-d326961673e3697">98</a></sup> were used as references. Multiple sequence alignments were performed using the MAFFT<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 88" title="Katoh, K., Kuma, K. I., Toh, H. & Miyata, T. MAFFT version 5: improvement in accuracy of multiple sequence alignment. Nucleic Acids Res. 33, 511–518 (2005)." href="/articles/s41477-019-0588-4#ref-CR88" id="ref-link-section-d326961673e3701">88</a></sup> program (<a href="https://mafft.cbrc.jp/alignment/software/">https://mafft.cbrc.jp/alignment/software/</a>). The maximum-likelihood phylogenetic trees were implemented with RAxML-HPC2 on XSEDE<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 101" title="Stamatakis, A., Hoover, P. & Rougemont, J. A rapid bootstrap algorithm for the RAxML Web servers. Syst. Biol. 57, 758–771 (2008)." href="/articles/s41477-019-0588-4#ref-CR101" id="ref-link-section-d326961673e3713">101</a></sup> through the CIPRES Science Gateway (v.3.3) (<a href="https://www.phylo.org/">https://www.phylo.org/</a>), estimating branch support values by bootstrap iterations with 1,000 replicates.</p><h3 class="c-article__sub-heading" id="Sec23">Gene-family expansion identification</h3><p>To understand gene-family expansion or contraction in <i>A. angustus</i> compared with that in 18 other green plants, the mean gene-family size was calculated for all gene families (excluding orphans and species-specific families). The number of genes per species for each family was transformed into a matrix of z-scores to centre and normalize the data. The first 100 families with the largest gene-family size in <i>A</i>. <i>angustus</i> were selected (Supplementary Fig. <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">55</a>). The clustering and visualization were performed using Genesis (v.3.0)<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 102" title="Sturn, A., Quackenbush, J. & Trajanoski, Z. Genesis: cluster analysis of microarray data. Bioinformatics 18, 207–208 (2002)." href="/articles/s41477-019-0588-4#ref-CR102" id="ref-link-section-d326961673e3744">102</a></sup>. The functional annotation of each family was predicted on the basis of sequence similarity to entries in the Pfam protein domain database, where more than 30% of proteins in the family share the same protein domain. Transposon-derived gene families were removed because the distribution of such families is likely to be a consequence of the gene models derived from a repeat-masked genome sequence and therefore may be artefactual<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 103" title="Martens, C., Vandepoele, K. & Van de Peer, Y. Whole-genome analysis reveals molecular innovations and evolutionary transitions in chromalveolate species. Proc. Natl Acad. Sci. USA 105, 3427–3432 (2008)." href="/articles/s41477-019-0588-4#ref-CR103" id="ref-link-section-d326961673e3749">103</a></sup>.</p><h3 class="c-article__sub-heading" id="Sec24">Tandem duplication definition</h3><p>Genes were defined as tandemly arrayed genes if they belonged to the same family, were located within 100 kb each other, and were separated by zero, one or fewer, five or fewer, or ten or fewer non-homologous intervening ‘spacer’ genes<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 104" title="Hanada, K. et al. Importance of lineage-specific expansion of plant tandem duplicates in the adaptive response to environmental stimuli. Plant Physiol. 148, 993–1003 (2008)." href="/articles/s41477-019-0588-4#ref-CR104" id="ref-link-section-d326961673e3761">104</a></sup>. Therefore, the four sets of tandem gene definitions were analysed.</p><h3 class="c-article__sub-heading" id="Sec25">HGT event identification</h3><p>In this study, we used two different strategies to identify candidates for <i>A. angustus</i>-specific and bryophyte-specific HGTs. For <i>A. angustus</i>-specific HGTs, we submitted 14,629 predicted coding genes of <i>A. angustus</i> to a BLASTP search against the NCBI protein database (<i>E</i>-value cutoff of 1 × 10<sup>−5</sup>) (Supplementary Note <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">7.1</a>). The proteins with the best BLAST hits in bacterial or fungal sequences were extracted. After sequences without support of transcript evidence were excluded, a series of parameters were used to filter the candidates (Supplementary Note <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">7.1</a>). For the bryophyte-specific HGT, we extracted gene families that are common to at least two of the three members of bryophytes (moss <i>P. patens</i>, liverwort <i>M. polymorpha</i> and hornwort <i>A. angustus</i>). To preliminarily determine whether these clusters are HGT candidates, we submitted the corresponding <i>A. angustus</i> members of each cluster to the NCBI protein database for BLASTP search and checked the taxonomy report of the top 1,000 BLAST hits (Supplementary Note <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM1">7.2</a>). The homologues of published HGTs in <i>P. patens</i><sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 51" title="Yue, J., Hu, X., Sun, H., Yang, Y. & Huang, J. Widespread impact of horizontal gene transfer on plant colonization of land. Nat. Commun. 3, 1152 (2012)." href="/articles/s41477-019-0588-4#ref-CR51" id="ref-link-section-d326961673e3812">51</a></sup> and <i>M. polymorpha</i><sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 15" title="Bowman, J. L. et al. Insights into land plant evolution garnered from the Marchantia polymorpha genome. Cell 171, 287–304 (2017)." href="/articles/s41477-019-0588-4#ref-CR15" id="ref-link-section-d326961673e3818">15</a></sup> were also investigated in the <i>A. angustus</i> genome. All candidate HGTs were subjected to phylogenetic analysis for verification. Synonymous codon-usage order values and GC contents of HGT and non-HGT genes were calculated by CodonO<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 105" title="Angellotti, M. C., Bhuiyan, S. B., Chen, G., Wan, X. & Wan, X. CodonO: codon usage bias analysis within and across genomes. Nucleic Acids Res. 35, W132–W136 (2007)." href="/articles/s41477-019-0588-4#ref-CR105" id="ref-link-section-d326961673e3825">105</a></sup>.</p><h3 class="c-article__sub-heading" id="Sec26">Reporting Summary</h3><p>Further information on experimental design is available in the <a data-track="click" data-track-label="link" data-track-action="supplementary material anchor" href="/articles/s41477-019-0588-4#MOESM2">Nature Research Reporting Summary</a> linked to this article.</p></div></div></section> </div> <div class="u-mt-32"> <section data-title="Data availability"><div class="c-article-section" id="data-availability-section"><h2 class="c-article-section__title js-section-title js-c-reading-companion-sections-item" id="data-availability">Data availability</h2><div class="c-article-section__content" id="data-availability-content"> <p>The <i>A. angustus</i> genome project has been deposited at the NCBI under the BioProject number <a href="https://www.ncbi.nlm.nih.gov/bioproject/?term=PRJNA543716">PRJNA543716</a>. The genome sequencing data were deposited in the Sequence Read Archive database under the accession number <a href="http://www.ncbi.nlm.nih.gov/sra?term=SRR9696346">SRR9696346</a>. The <i>A. angustus</i> transcriptome project has been deposited at the NCBI under BioProject <a href="https://www.ncbi.nlm.nih.gov/bioproject/?term=PRJNA543724">PRJNA543724</a>. The transcriptome sequencing data were deposited in the Sequence Read Archive database under the accession number <a href="http://www.ncbi.nlm.nih.gov/sra?term=SRR9662965">SRR9662965</a>. The assembled genome sequences, gene models and miRNA data are available via DRYAD (<a href="https://doi.org/10.5061/dryad.msbcc2ftv">https://doi.org/10.5061/dryad.msbcc2ftv</a>). 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R. Crane, S. Ge, D.-Y. Hong, J.-L. Huang, J.-J. Qin, Y.-L. Qiu, J.-C. Villarreal, T. Wan and X.-Q. Wang for useful advice and discussions and L. Zhang for providing plant pictures. We dedicate the paper to Yang Zhong in memory of his support and valuable suggestions on this project. This work was supported by Sino–Africa Joint Research Center, Chinese Academy of Sciences, CAS International Research and Education Development Program (SAJC201613), the Strategic Priority Research Program of the Chinese Academy of Sciences (XDB31000000 and XDA19050103), National Natural Science Foundation of China (NNSF 31590822), Shenzhen Fairy Lake Botanical Garden, State Key Laboratory of Systematic and Evolutionary Botany, Institute of Botany, Chinese Academy of Sciences, and Key Laboratory of National Forestry and Grassland Administration for Orchid Conservation and Utilization at College of Landscape Architecture, Fujian Agriculture and Forestry University. Y.V.d.P. acknowledges support from the European Union Seventh Framework Programme (FP7/2007-2013) under European Research Council Advanced Grant Agreement 322739–DOUBLEUP.</p></div></div></section><section aria-labelledby="author-information" data-title="Author information"><div class="c-article-section" id="author-information-section"><h2 class="c-article-section__title js-section-title js-c-reading-companion-sections-item" id="author-information">Author information</h2><div class="c-article-section__content" id="author-information-content"><span class="c-article-author-information__subtitle u-visually-hidden" id="author-notes">Author notes</span><ol class="c-article-author-information__list"><li class="c-article-author-information__item" id="na1"><p>These authors contributed equally: Jian Zhang, Xin-Xing Fu, Rui-Qi Li, Xiang Zhao, Yang Liu, Ming-He Li, Arthur Zwaenepoel.</p></li><li class="c-article-author-information__item" id="na2"><p>These authors jointly supervised this work: Shou-Zhou Zhang, Yves Van de Peer, Zhong-Jian Liu, Zhi-Duan Chen.</p></li></ol><h3 class="c-article__sub-heading" id="affiliations">Authors and Affiliations</h3><ol class="c-article-author-affiliation__list"><li id="Aff1"><p class="c-article-author-affiliation__address">State Key Laboratory of Systematic and Evolutionary Botany, Institute of Botany, Chinese Academy of Sciences, Beijing, China</p><p class="c-article-author-affiliation__authors-list">Jian Zhang, Xin-Xing Fu, Rui-Qi Li, Qing-Hua Wang, Yu Jia, Mei-Zhi Wang, Yuan-Nian Jiao, Ya-Long Guo, Hong-Zhi Kong, An-Ming Lu & Zhi-Duan Chen</p></li><li id="Aff2"><p class="c-article-author-affiliation__address">University of Chinese Academy of Sciences, Beijing, China</p><p class="c-article-author-affiliation__authors-list">Xin-Xing Fu</p></li><li id="Aff3"><p class="c-article-author-affiliation__address">PubBio-Tech Services Corporation, Wuhan, China</p><p class="c-article-author-affiliation__authors-list">Xiang Zhao & Zhi-Wen Wang</p></li><li id="Aff4"><p class="c-article-author-affiliation__address">Key Laboratory of Southern Subtropical Plant Diversity, Fairy Lake Botanical Garden, Shenzhen & Chinese Academy of Science, Shenzhen, China</p><p class="c-article-author-affiliation__authors-list">Yang Liu, Yi-Ying Liao, Shan-Shan Dong, Jian-Fen Yang & Shou-Zhou Zhang</p></li><li id="Aff5"><p class="c-article-author-affiliation__address">BGI-Shenzhen, Shenzhen, China</p><p class="c-article-author-affiliation__authors-list">Yang Liu & Huan-Ming Yang</p></li><li id="Aff6"><p class="c-article-author-affiliation__address">Key Laboratory of National Forestry and Grassland Administration for Orchid Conservation and Utilization at College of Landscape Architecture, Fujian Agriculture and Forestry University, Fuzhou, China</p><p class="c-article-author-affiliation__authors-list">Ming-He Li, Jie-Yu Wang, Guo-Qiang Zhang & Zhong-Jian Liu</p></li><li id="Aff7"><p class="c-article-author-affiliation__address">Department of Plant Biotechnology and Bioinformatics, Ghent University, Ghent, Belgium</p><p class="c-article-author-affiliation__authors-list">Arthur Zwaenepoel & Yves Van de Peer</p></li><li id="Aff8"><p class="c-article-author-affiliation__address">VIB Center for Plant Systems Biology, Ghent, Belgium</p><p class="c-article-author-affiliation__authors-list">Arthur Zwaenepoel & Yves Van de Peer</p></li><li id="Aff9"><p class="c-article-author-affiliation__address">Department of Biology, Huck Institutes of the Life Sciences, Pennsylvania State University, University Park, PA, USA</p><p class="c-article-author-affiliation__authors-list">Hong Ma</p></li><li id="Aff10"><p class="c-article-author-affiliation__address">Department of Ecology and Evolutionary Biology, University of Connecticut, Storrs, CT, USA</p><p class="c-article-author-affiliation__authors-list">Bernard Goffinet</p></li><li id="Aff11"><p class="c-article-author-affiliation__address">Key Laboratory for Plant Diversity and Biogeography of East Asia, Kunming Institute of Botany, Chinese Academy of Sciences, Kunming, China</p><p class="c-article-author-affiliation__authors-list">Yan-Long Guan</p></li><li id="Aff12"><p class="c-article-author-affiliation__address">Center for Plant Diversity and Systematics, Institute of Botany, Jiangsu Province and Chinese Academy of Sciences, Nanjing, China</p><p class="c-article-author-affiliation__authors-list">Jia-Yu Xue</p></li><li id="Aff13"><p class="c-article-author-affiliation__address">Sino–Africa Joint Research Center, Chinese Academy of Sciences, Wuhan, China</p><p class="c-article-author-affiliation__authors-list">Yi-Ying Liao, Qing-Feng Wang & Zhi-Duan Chen</p></li><li id="Aff14"><p class="c-article-author-affiliation__address">College of Forestry and Landscape Architecture, South China Agricultural University, Guangzhou, China</p><p class="c-article-author-affiliation__authors-list">Jie-Yu Wang & Zhong-Jian Liu</p></li><li id="Aff15"><p class="c-article-author-affiliation__address">Center for Microbial Ecology and Genomics, Department of Biochemistry, Genetics and Microbiology, Pretoria, South Africa</p><p class="c-article-author-affiliation__authors-list">Yves Van de Peer</p></li><li id="Aff16"><p class="c-article-author-affiliation__address">College of Horticulture, Nanjing Agricultural University, Nanjing, China</p><p class="c-article-author-affiliation__authors-list">Yves Van de Peer</p></li><li id="Aff17"><p class="c-article-author-affiliation__address">Fujian Colleges and Universities Engineering Research Institute of Conservation and Utilization of Natural Bioresources, College of Forestry, Fujian Agriculture and Forestry University, Fuzhou, China</p><p class="c-article-author-affiliation__authors-list">Zhong-Jian Liu</p></li></ol><div class="u-js-hide u-hide-print" data-test="author-info"><span class="c-article__sub-heading">Authors</span><ol class="c-article-authors-search u-list-reset"><li 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Guan. conducted DAPI staining analysis; J.-Y.X. conducted codon-usage bias analysis; M.-H.L., G.-Q.Z. and J.-Y.W. conducted transcriptome sequencing and analysis; S.-S.D. and Y.L. conducted the RNA-editing-site analysis in organellar genomes; H.M., Q.-F.W., B.G., Y.J., Y.-N.J., Y.-L.Guo, H.-Z.K., A.-M.L. and H.-M.Y. contributed substantially to revisions. All authors commented on the manuscript.</p><h3 class="c-article__sub-heading" id="corresponding-author">Corresponding authors</h3><p id="corresponding-author-list">Correspondence to <a id="corresp-c1" href="mailto:shouzhouz@126.com">Shou-Zhou Zhang</a>, <a id="corresp-c2" href="mailto:yves.vandepeer@psb.vib-ugent.be">Yves Van de Peer</a>, <a id="corresp-c3" href="mailto:zjliu@fafu.edu.cn">Zhong-Jian Liu</a> or <a id="corresp-c4" href="mailto:zhiduan@ibcas.ac.cn">Zhi-Duan Chen</a>.</p></div></div></section><section data-title="Ethics declarations"><div class="c-article-section" id="ethics-section"><h2 class="c-article-section__title js-section-title js-c-reading-companion-sections-item" id="ethics">Ethics declarations</h2><div class="c-article-section__content" id="ethics-content"> <h3 class="c-article__sub-heading" id="FPar1">Competing interests</h3> <p>The authors declare no competing financial interests.</p> </div></div></section><section data-title="Additional information"><div class="c-article-section" id="additional-information-section"><h2 class="c-article-section__title js-section-title js-c-reading-companion-sections-item" id="additional-information">Additional information</h2><div class="c-article-section__content" id="additional-information-content"><p><b>Peer review information</b> Nature Plants thanks Burkhard Becker and the other, anonymous, reviewers for their contribution to the peer review of this work.</p><p><b>Publisher’s note</b> Springer Nature remains neutral with regard to jurisdictional claims in published maps and institutional affiliations.</p></div></div></section><section data-title="Supplementary information"><div class="c-article-section" id="Sec27-section"><h2 class="c-article-section__title js-section-title js-c-reading-companion-sections-item" id="Sec27">Supplementary information</h2><div class="c-article-section__content" id="Sec27-content"><div data-test="supplementary-info"><div id="figshareContainer" class="c-article-figshare-container" data-test="figshare-container"></div><div class="c-article-supplementary__item" data-test="supp-item" id="MOESM1"><h3 class="c-article-supplementary__title u-h3"><a class="print-link" data-track="click" data-track-action="view supplementary info" data-test="supp-info-link" data-track-label="supplementary information" href="https://static-content.springer.com/esm/art%3A10.1038%2Fs41477-019-0588-4/MediaObjects/41477_2019_588_MOESM1_ESM.pdf" data-supp-info-image="">Supplementary Information</a></h3><div class="c-article-supplementary__description" data-component="thumbnail-container"><p>Supplementary Notes 1–7, Figs. 1–87 and Tables 1–20.</p></div></div><div class="c-article-supplementary__item" data-test="supp-item" id="MOESM2"><h3 class="c-article-supplementary__title u-h3"><a class="print-link" data-track="click" data-track-action="view supplementary info" data-test="supp-info-link" data-track-label="reporting summary" href="https://static-content.springer.com/esm/art%3A10.1038%2Fs41477-019-0588-4/MediaObjects/41477_2019_588_MOESM2_ESM.pdf" data-supp-info-image="">Reporting Summary</a></h3></div><div class="c-article-supplementary__item" data-test="supp-item" id="MOESM3"><h3 class="c-article-supplementary__title u-h3"><a class="print-link" data-track="click" data-track-action="view supplementary info" data-test="supp-info-link" data-track-label="supplementary tables" href="https://static-content.springer.com/esm/art%3A10.1038%2Fs41477-019-0588-4/MediaObjects/41477_2019_588_MOESM3_ESM.xlsx" data-supp-info-image="">Supplementary Tables</a></h3><div class="c-article-supplementary__description" data-component="thumbnail-container"><p>Supplementary Tables 21–32.</p></div></div></div></div></div></section><section data-title="Rights and permissions"><div class="c-article-section" id="rightslink-section"><h2 class="c-article-section__title js-section-title js-c-reading-companion-sections-item" id="rightslink">Rights and permissions</h2><div class="c-article-section__content" id="rightslink-content"> <p><b>Open Access</b> This article is licensed under a Creative Commons Attribution 4.0 International License, which permits use, sharing, adaptation, distribution and reproduction in any medium or format, as long as you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons license, and indicate if changes were made. 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id="citeas">Cite this article</h3><p class="c-bibliographic-information__citation">Zhang, J., Fu, XX., Li, RQ. <i>et al.</i> The hornwort genome and early land plant evolution. <i>Nat. Plants</i> <b>6</b>, 107–118 (2020). https://doi.org/10.1038/s41477-019-0588-4</p><p class="c-bibliographic-information__download-citation u-hide-print"><a data-test="citation-link" data-track="click" data-track-action="download article citation" data-track-label="link" data-track-external="" rel="nofollow" href="https://citation-needed.springer.com/v2/references/10.1038/s41477-019-0588-4?format=refman&flavour=citation">Download citation<svg width="16" height="16" focusable="false" role="img" aria-hidden="true" class="u-icon"><use xmlns:xlink="http://www.w3.org/1999/xlink" xlink:href="#icon-eds-i-download-medium"></use></svg></a></p><ul class="c-bibliographic-information__list" data-test="publication-history"><li class="c-bibliographic-information__list-item"><p>Received<span class="u-hide">: </span><span class="c-bibliographic-information__value"><time datetime="2019-06-12">12 June 2019</time></span></p></li><li class="c-bibliographic-information__list-item"><p>Accepted<span 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