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Here we report the presence of lianas in the Zhangpu flora based on exquisitely preserved fossil flowers of Ayenia.<\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Methods<\/jats:title>\n                    <jats:p>Two fossil flowers, preserved in amber, were investigated with light microscopy and X-ray tomography, and are described herein as Ayenia zhangpuensis sp. nov. and Ayenia sp.<\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Key Results<\/jats:title>\n                    <jats:p>The flower of Ayenia zhangpuensis is bisexual and pentamerous. Its sepals are ovate, the petals are clawed, with an inner ridge on their hood delimiting a fossa, together forming a lantern-like corolla around the androecium and gynoecium. The androecium forms a campanulate staminal tube; the gynoecium is syncarpous with a superior ovary. The other specimen, treated informally as Ayenia sp., differs from A. zhangpuensis in having more incurved and elongate sepals. The two fossil flowers closely resemble extant flowers of the Ayenia section Vahihara (former Byttneria section Vahihara), almost all species of which are unarmed lianas thriving in humid tropical conditions.<\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Conclusions<\/jats:title>\n                    <jats:p>The striking similarities between fossil and extant flowers of the Ayenia section Vahihara suggest a morphological stasis in floral structure within a tropical liana lineage since at least the mid-Miocene. Together with other climbing plants known from the Zhangpu flora, our study shows that this Miocene rainforest is comparable to modern Southeast Asian rainforests in both floristic composition and inferred forest structure.<\/jats:p>\n                  <\/jats:sec>","DOI":"10.1093\/aob\/mcag220","type":"journal-article","created":{"date-parts":[[2026,7,16]],"date-time":"2026-07-16T11:50:24Z","timestamp":1784202624000},"source":"Crossref","is-referenced-by-count":1,"title":["Fossil flowers of\n                    <i>Ayenia<\/i>\n                    (Malvaceae) preserved in amber: evidence for lianas in Miocene rainforest of southeast China"],"prefix":"10.1093","author":[{"given":"Xieting","family":"Wu","sequence":"first","affiliation":[{"name":"State Key Laboratory of Palaeobiology and Stratigraphy, Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences , Nanjing 210008 ,","place":["China"]},{"name":"Department of Botany and Biodiversity Research, University of Vienna , Vienna 1030 ,","place":["Austria"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Simon","family":"Beurel","sequence":"additional","affiliation":[{"name":"Museum f\u00fcr Naturkunde, Leibniz Institute for Evolution and Biodiversity Science , Berlin 10115 ,","place":["Germany"]},{"name":"Freie Universit\u00e4t Berlin, Institute of Biology , Berlin 14195 ,","place":["Germany"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-9472-1615","authenticated-orcid":false,"given":"Julien B","family":"Bachelier","sequence":"additional","affiliation":[{"name":"Freie Universit\u00e4t Berlin, Institute of Biology , Berlin 14195 ,","place":["Germany"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Maria","family":"von Balthazar","sequence":"additional","affiliation":[{"name":"Department of Botany and Biodiversity Research, University of Vienna , Vienna 1030 ,","place":["Austria"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"J\u00f6rg U","family":"Hammel","sequence":"additional","affiliation":[{"name":"Institute of Materials Physics, Helmholtz-Zentrum Hereon , Geesthacht 21502 ,","place":["Germany"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Eva-Maria","family":"Sadowski","sequence":"additional","affiliation":[{"name":"Museum f\u00fcr Naturkunde, Leibniz Institute for Evolution and Biodiversity Science , Berlin 10115 ,","place":["Germany"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-3374-6637","authenticated-orcid":false,"given":"Gongle","family":"Shi","sequence":"additional","affiliation":[{"name":"State Key Laboratory of Palaeobiology and Stratigraphy, Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences , Nanjing 210008 ,","place":["China"]},{"name":"University of Chinese Academy of Sciences, Nanjing , Nanjing 211135 ,","place":["China"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"286","published-online":{"date-parts":[[2026,7,17]]},"reference":[{"key":"2026081003091337500_mcag220-B1","doi-asserted-by":"publisher","first-page":"77","DOI":"10.3140\/bull.geosci.1145","article-title":"Paleocene and Eocene floristic and climatic change in Russia and Northern Kazakhstan","volume":"85","author":"Akhmetiev","year":"2010","journal-title":"Bulletin of Geosciences"},{"key":"2026081003091337500_mcag220-B2","first-page":"104","article-title":"Estructura y ontogenia de los nectarios foliares del g\u00e9nero Byttneria (Sterculiaceae)","volume":"17","author":"Arbo","year":"1972","journal-title":"Darwiniana"},{"key":"2026081003091337500_mcag220-B3","doi-asserted-by":"publisher","first-page":"267","DOI":"10.1111\/j.1095-8339.1999.tb00505.x","article-title":"Support for an expanded family concept of Malvaceae within a recircumscribed order Malvales: a combined analysis of plastid atpB and rbcL DNA sequences","volume":"129","author":"Bayer","year":"1999","journal-title":"Botanical Journal of the Linnean Society"},{"key":"2026081003091337500_mcag220-B4","doi-asserted-by":"crossref","first-page":"225","DOI":"10.1007\/978-3-662-07255-4_28","volume-title":"Flowering plants: dicotyledons. 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