Early Permian fossil fruit reshapes angiosperm history

Early Permian fossil fruit reshapes angiosperm history | Research Square window.SnipcartSettings = { analytics: { enabled: false } }; (function() { var accessVector = localStorage.getItem('access_vector') || ''; window.dataLayer = window.dataLayer || []; if (accessVector) { window.dataLayer.push({ user: { profile: { profileInfo: { snid: accessVector } } } }); } })(); (function(w,d,s,l,i){w[l]=w[l]||[];w[l].push({'gtm.start':new Date().getTime(),event:'gtm.js'});var f=d.getElementsByTagName(s)[0],j=d.createElement(s),dl=l!='dataLayer'?'&l='+l:'';j.async=true;j.src='https://www.googletagmanager.com/gtm.js?id='+i+dl;f.parentNode.insertBefore(j,f);})(window,document,'script','dataLayer','GTM-K279D39R'); Browse Preprints In Review Journals COVID-19 Preprints AJE Video Bytes Research Tools Research Promotion AJE Professional Editing AJE Rubriq About Preprint Platform In Review Editorial Policies Our Team Advisory Board Help Center Sign In Submit a Preprint Cite Share Download PDF Systematic Review Early Permian fossil fruit reshapes angiosperm history Xin Wang, Weijia Huang, Qiang Fu, Yinggang Lei This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-7244136/v1 This work is licensed under a CC BY 4.0 License Status: Posted Version 1 posted You are reading this latest preprint version Abstract Angiosperms are the most important plant group for humans in the current earth ecosystem 1 . However, the origin and early history of angiosperms remain elusive and controversial 2 – 9 . According to the over60-year-old stereotype in botany, angiosperms cannot be older than the Cretaceous 10 , 11 . However, this thinking is now facing increasing challenges in terms of fossil evidence 6 , 12 – 17 and molecular estimates 7 – 9 . Apparently, serious and independent studies on new early fossil angiosperms are urgently needed to resolve this academic confrontation. In recent years, a renewed wave of interest in Permian fossil plants in Henan’s Cathaysian flora has been rekindled by the latest palaeobotanical progress 15 , 16 . During a recent field excursion in May 2025, we collected a new fossil organ from the Lower Shihezi (formerly Shihhotse)Formation (Lower Permian) of Dengfeng, Henan, China. This new fossil organ is interpreted as a large fruit and named Dengfengfructus maxima gen. et sp. nov. The seed enclosed in the fruit has a peripheral three-layered seed coat, which distinguishes the seed itself from an otherwise possible nucellus in a seed. The good preservation allows the cellular details in the seed coat and seed content to be revealed. Unlike contemporary Taiyuanostachya 16 and Yuzhoua 15 , Dengfengfructus has a large size and a thick pericarp. This new fossil organ apparently updates and enhances the current understanding of angiosperms and their diversity in the Permian. The history of angiosperms can be dated back to the Early Permian (Palaeozoic). The former stereotype that angiosperms are restricted to the Cretaceous and later ages falls apart in front of the currently available palaeobotanical progress and molecular estimates. Obviously, we are facing a wave of botanical theories in the near future. angiosperm fruit Permian Palaeozoic 294.4 Ma Figures Figure 1 Figure 2 Figure 3 Main text Although angiosperms account for most of the species diversity of plants on Earth 1 , little is known about the origin and early history of angiosperms. The publishing of Taiyuanostachya 16 and Yuzhoua 15 as Early Permian angiosperms opened a new era of angiosperm study. However, there is a confrontation between some palaeobotanists who insisted on the Early Cretaceous origin for angiosperms 10,11 and the phylogenomic scientists who found a pre-Cretaceous origin for angiosperms 7-9 as well as palaeobotanists who unveiled pre-Cretaceous angiosperms 12-17 . The only reliable way to elucidate the early history of angiosperms is to study more fossils of plants. The Yuzhou Flora is typical of the Lower Shihezi (formerly Shihhotse)Formation, Lower Permian in Henan, China 18 , and is one of the best studied parts of the Cathayasian Flora 19-21 . Previous studies indicate that the Yuzhou Flora comprises 307 species in 111 genera of Sphenophyllales, Equisetales, Filicales, Lepidodendrales, Noeggerathiales, Peltaspermales, Pteridospermophytes, Cycadales, Dicranohpyllopsida, Ginkgophyta, Cordaitopsida, Coniferopsida, Gigantopteridales, and plants incertae sedis 18 . The age of this flora has been dated as the Kungurian–Roadian Asselian 22 , namely, 294.4–295.1 Ma 23 . The specimen was collected from an outcrop of the Lower Shihezi (formerly Shihhotse)Formation near Wangyao village, Gaocheng town, Dengfeng city, Henan Province, China (113.155925°E, 34.341893°N; Fig. 1) by Y.L. in May, 2025. The fossil was exposed on the surface naturally, requiring little trimming or dégagement. The morphology was photographed using a Nikon D800. The details of the specimen were photographed via a Zeiss Zoom V16 stereomicroscope. All the figures were organized for publication in Photoshop 7.0 software. Angiosperms Order incertae sedis Family incertae sedis Dengfengfructus gen. nov. Type species : Dengfengfructus maxima gen. et sp. nov. Diagnosis : Drupe oval, enclosing a single seed. Seed comprising a three-layered seed coat and seed content. Dengfengfructus maxima gen. et sp. nov. (Figures 2–3) Diagnosis : The same as that of the genus, for the time being. Description: The fruit is preserved as a compression, oval–shaped, 17.6 mm long and 14 mm wide (Fig. 2a). The fruit comprises a seed and a surrounding pericarp (Fig. 2a). The thickness of the pericarp varies from 2 to 3.3 mm (Fig. 2a). The pericarp appears homogeneous in structure, with a basal vascular bundle and secluded tip, with cellular details (Figs. 2a-c, 3a). The seed is approximately 10 mm in diameter and 12 mm tall, oval–shaped, and surrounded by a three-layered seed coat containing a seed content that is in turn fully enclosed by the pericarp (Figs. 2a, d-e, 3b-d). The three layers of the seed coat are 470 μm, 390 μm, and 240 μm thick, respectively, from inner to outer (Fig. 2e). The cells in the seed coat have conspicuous angular outlines, 82~210 μm × 145~424 μm, with little differentiation among the layers (Fig. 2e). In contrast, the seed content is homogeneous, including cells with inconspicuous outlines of 75~118 μm × 101~193 μm (Fig. 2f). The fruit is associated with the leaves of Lobatannularia spatulata (He) Yang (Fig. S1a). Etymology : Dengfengfructus is for a fruit from the city of Dengfeng in Henan, China, where the fossil is from. The specific epithet maxima is for the large size of the fruit. Holotype specimen: PB207444. Depository: Nanjing Institute of Geology and Palaeontology, Nanjing, China. Type locality : Wangyao village, Gaocheng town, Dengfeng city, Henan Province, China (113.155925°E, 34.341893°N). Horizon : the Lower Shihezi (formerly Shihhotse)Formation, Permian (294.4-295.1 Ma) 23 . Remarks: The age of our fossil is supported by its associated leaves, Lobatannularia spatulata (He) Yang 2006, preserved on the same specimen (Fig. S1a). Lobatannularia spatulata (He) Yang is frequently seen in the Yuzhou Flora and has been well studied in previous works 18 . The co-occurrence of these two fossils on the same piece of specimen, on one hand, is suggestive of the abundant occurrence of plant fossils in the strata; on the other hand, the repeated occurrences of Lobatannularia spatulata (He) Yang in various studies pin down the Early Permian age for Dengfengfructus 18 . Two recent thorough stratigraphic surveys 22,23 suggested a Kungurian–Roadian Asselian age (294.4-295.1 Ma) for our fossil plant. The single seed of Dengfengfructus is fully enclosed by a pericarp, satisfying the original criterion for angiosperms given by Paul Hermann in 1690 24 . This is the main reason we interpret Dengfengfructus as an angiosperm fruit. However, considering Palaeozoic angiosperms are still rare and that seeds are most likely present in the Early Permian, we prefer to exclude all alternative interpretations before proceeding further. Theoval general morphology of Dengfengfructus distinguishes itself from all known vegetative parts of plants. Among the reproductive organs of plants, there are no male organs of similar size or organization. Therefore, we do not consider any male reproductive organs in the following discussion. The most plausible alternatives for Dengfengfructus are a female reproductive organ or its part (namely, a fruit, an ovule, or a seed). Since ovules and seeds were observed as early as the Late Devonian, we must consider these alternatives seriously for our Permian fossil organ. The morphology and organization of Palaeozoic ovules/seeds are well known 25 . A mature Palaeozoic seed usually has a seed content surrounded by a three-layered seed coat (including sarcotesta, sclerotesta, and endotesta) 25 . In contrast to seed content, which is unlikely to be preserved in fossils, a seed coat is the part that has the greatest potential to be fossilized in a seed. This generalization contrasts with our observations: the peripheral surrounding layer of Dengfengfructus is homogeneous and shows no trace of stratification (Fig. 2a); thus, it is distinct from any known seed coat. Apparently, Dengfengfructus cannot be compared to any mature seeds of any plant. How about an immature seed? An immature seed usually shows a smooth and gradual transition to an ovule in morphology and organization. An ovule usually includes a nucellus and protective integuments. Integuments (corresponding to the seed coat in a mature seed) partially surround the nucellus and may appear homogeneous when immature or may demonstrate stratification when mature, whereas the nucellus corresponds to the seed content in a mature seed and usually demonstrates no stratification. This again contradicts our observation: the central oval body in Dengfengfructus apparently has a three-layered peripheral part and a central homogeneous content (Figs. 2a, d-f, 3b-d). By the way, the seeds of interesting plants in Bennetitales have their characteristic micropylar tubes, excluding themselves from our consideration. The above contrasts and comparisons exclude the possibility of Dengfengfructus being either a seed or an ovule, leaving only one alternative for Dengfengfructus : fruit. Although a Permian angiosperm is unacceptable for many, as it is far older than the widely accepted age for angiosperms (the Early Cretaceous 10,11 ), recent phylogenomic studies and palaeobotanical studies of Taiyuanostachya 16 and Yuzhoua 15 have undermined the credibility of the Cretaceous origin for angiosperms 10,11 , making the angiospermous affinity of Dengfengfructus less surprising. According to the traditional thought dating back to the age of Arber and Parkin (1907), ancestral carpels are conduplicate carpels with marginal ovules 26 . However, this expectation is not honored by fossil evidence. For example, although Dilcher and Crane 27 appeared to support traditional thinking via their fossil evidence of Arachaeanthus , subsequent careful examination indicated that the seeds/ovules in Archaeanthus are actually borne at the dorsal margin of the fruit/carpel, thus refuting the traditional thinking 28 . In our case, the basally fixed seed in Dengfengfructus is apparently at odds with the widely expected marginally-positioned seeds, further reducing the credibility of the traditional thinking and lending support to the recently advanced Unifying Theory 29,30 , which allowed various positions of a seed in a fruit. Notably, even studies on carpels of the assumed ancestral living Magnoliales ( Michelia 31 and Illicium 32 ) have rejected the traditional thinking. On the basis of currently available data, the traditional thinking of angiosperm evolution, although widely accepted and taught in classrooms, is highly flawed and should be discarded. A caveat of the current conclusion that Dengfengfructus is an angiosperm, the pollination process of which is unknown. Since Tomlinson and Takaso 33 have distinguished angiosperms from gymnosperms by the ovule-enclosing status of a plant at the time of pollination, namely, ovule-enclosing occurs before pollination in angiosperms. This reduces the validity of our above judgement, but it does not reduce the evolutionary significance of Dengfengfructus. There are only two possible scenarios in the future. 1) If a future study proves that ovule enclosure occurs before pollination in Dengfengfructus , then Dengfengfructus will be confirmed as a bona fide Palaeozoic angiosperm. In this case, the existence of angiosperms in the Early Permian will be verified and consolidated, and angiosperm diversity in the Permian (Palaeozoic) is further underscored, especially when Taiyuanostachya 16 and Yuzhoua 15 are taken into consideration. 2) However, if a future study proves that ovule/seed enclosure in Dengfengfructus occurs after pollination, then Dengfengfructus should be placed among gymnosperms. If Dengfengfructus were kicked out of angiosperms, it would not affect the Permian origin of angiosperms: Taiyuanostachya has not only demonstrates not only angiospermy (enclosed seed) but also angio-ovuly (enclosed ovule), thus having pushed the origin time of angiosperms into the EarlyPermian 16 . This alternative placement of Dengfengfructus means that Dengfengfructus , although a gymnosperm, has developed a seed-bearing structure that is hard to distinguish from an angiosperm fruit. This would reduce the former huge gap between angiosperms and gymnosperms, underscore the paper-thin and blurry boundary between angiosperms and gymnosperms, and corroborate the theoretically predicted smooth transition between angiosperms and gymnosperms. The timing order of two independent developmental events, ovule-enclosing and pollination, matters in terms of the taxonomy of seed plants and is the key feature distinguishing gymnosperms and angiosperms. During the long-term evolution of seed plants, some of them may arbitrarily sport and develop earlier ovule closure. This initially randomly occurring but later canalized and fixed feature apparently agrees with the general pattern of organism evolution, namely, offspring development conditioning (ODC) 34 . If such a feature benefits the plants in terms of proliferation, it is very likely to be canalized and carried on by their offspring. This developmental pattern may be heritable in plants. The end product of such a process is the debut of angiosperms in geological history. Conclusions Dengfengfructus maxima gen. et sp. nov. is a new Permian angiosperm fruit. Its Early Permian (294.4–295.1 Ma) age confirms the existence of angiosperms in the late Palaeozoic. The margin-independent position of seed in Dengfengfructus implies that the traditional thinking of angiosperms, which has been refuted by previous studies, is not favoured by our latest fossil evidence. The doubts cast over traditional thinking by various studies imply that we are facing a new wave of innovations in botanical theories, especially those concerned with angiosperm origin and evolution. Abbreviations SEM (scanning electron microscope) Declarations Ethics approval and consent to participate Not applicable Consent for publication Not applicable Funding The National Key Research and Development Program of China (2024YFF0807601). Authors' contributions WH and YL collected the samples. XW initiated the study and drafted the manuscript. QF, WH, YL and XW analysed the data and modified and finalized the manuscript. Acknowledgements We appreciate Ms. Jingjing Tang for assisting with the photography, and Mr. Yan Fang for assisting with SEM observation. Data availability All the data analysed during this study are included in this article. Competing interests The authors declare that they have no competing interests. References Crepet WL, Niklas KJ. 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The Dawn Angiosperms. Volume 407, 2nd ed. Springer; 2018. Wang X. The Dawn Angiosperms. Volume 236. Springer; 2010. Zhang X, Liu W, Wang X. How the ovules get enclosed in magnoliaceous carpels. PLoS ONE. 2017;12:e0174955. https://doi.org/10.1371/journal.pone.0174955 . Zhang X. Floral ontogeny of Illicium lanceolatum (Schisandraceae) and its implications on carpel homology. Phytotaxa. 2019;416:200–10. https://doi.org/10.11646/phytotaxa.416.3.1 . Tomlinson PB, Takaso T. Seed cone structure in conifers in relation to development and pollination: a biological approach. Can J Bot. 2002;80:1250–73. Fu Q, Liu J, Wang X. Offspring development conditioning (ODC): A universal evolutionary trend in sexual reproduction of organisms. J Northwest Univ (Natural Sci Edition). 2021;51:163–72. https://doi.org/10.16152/j.cnki.xdxbzr.2021-01-018 . Tables Table 1 Comparison among contemporaneous Dengfengfructus, Protofructus, Novofructus, Taiyuanostachya , and Yuzhoua . Dengfengfructus Protofructus Novofructus Taiyuanostachya Yuzhoua Preservation Compression Compression Compression Compression Compression Fossil locality Dengfeng, Henan Xinmi, Henan Xinmi, Henan Taiyuan, Shanxi Yuzhou, Henan Age Early Permian Early Permian Early Permian Early Permian Early Permian Horizon Lower Shihezi (formerly Shihhotse)Fm. Shape of organ oval elongated elongated cylindrical bulging Organization of organ sole fruit sole fruit sole fruit infructescence sole fruit Length of organ 17.6 mm 43 mm 36 mm 43 mm 40 mm Ovules per fruit 1 8+ 2+ 1 3 Ovule position basal on a long placenta along fruit margin basal in fruit, on infructescence axis on side ovarian wall Number of specimens 1 1 1 6 6 Additional Declarations No competing interests reported. Supplementary Files FigureS1.tif Figure S1 Dengfengfructus maxima gen. et sp. nov. and associated leaves of Lobatannularia spatulata (He) Yang 2006. 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Also discoverable on Platform About Our Team In Review Editorial Policies Advisory Board Help Center Resources Author Services Accessibility API Access RSS feed Manage Cookie Preferences © Research Square 2026 | ISSN 2693-5015 (online) Privacy Policy Terms of Service Do Not Sell My Personal Information {"props":{"pageProps":{"initialData":{"identity":"rs-7244136","acceptedTermsAndConditions":true,"allowDirectSubmit":true,"archivedVersions":[],"articleType":"Systematic Review","associatedPublications":[],"authors":[{"id":510539321,"identity":"9d0eaf0d-6b05-4120-b02a-a455a58654de","order_by":0,"name":"Xin Wang","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAA4klEQVRIiWNgGAWjYDACZgglB2MQr8WYBC1QkNhAtFKD47zHJH7uqE3fzs5+7cGPPwzy/A3Mzx7g0yLZzJcm2XvmeO7OZp5yw942BsMZB9jMDfBp4WfmMbvB23Ysd8NhnjQJ3gYGxg0MPGwS+LSwAbXc/Nt2LN0AqEXyzx8Ge4JaQLbc5m2rSTA4zH5MmoeNIZGgFslmHvPfsm0HDIEOYzeWbZNInnGYzQyvFoPzZ4wN37bVyRucP/7s4Zs/Nrb97c3P8GqBgsNAzGMGJCQYiI3TOiBmf0ac2lEwCkbBKBhxAAAKOUEOi5mPbAAAAABJRU5ErkJggg==","orcid":"","institution":"Chinese Academy of Sciences","correspondingAuthor":true,"prefix":"","firstName":"Xin","middleName":"","lastName":"Wang","suffix":""},{"id":510539322,"identity":"701f92a7-6509-4260-b52f-ba27f44bdcaf","order_by":1,"name":"Weijia Huang","email":"","orcid":"","institution":"Blue Miracle Museum Science Research Studio","correspondingAuthor":false,"prefix":"","firstName":"Weijia","middleName":"","lastName":"Huang","suffix":""},{"id":510539323,"identity":"7980678a-2762-4b27-9abd-0afaa9d5f7db","order_by":2,"name":"Qiang Fu","email":"","orcid":"","institution":"Chinese Academy of Sciences","correspondingAuthor":false,"prefix":"","firstName":"Qiang","middleName":"","lastName":"Fu","suffix":""},{"id":510539324,"identity":"b4c822a4-811a-43fa-afad-c6125015f262","order_by":3,"name":"Yinggang Lei","email":"","orcid":"","institution":"Henan University of Science and Technology","correspondingAuthor":false,"prefix":"","firstName":"Yinggang","middleName":"","lastName":"Lei","suffix":""}],"badges":[],"createdAt":"2025-07-29 14:08:19","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-7244136/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-7244136/v1","draftVersion":[],"editorialEvents":[],"editorialNote":"","failedWorkflow":false,"files":[{"id":90868743,"identity":"ab489a3f-b19c-4fc8-aec9-9734aece892b","added_by":"auto","created_at":"2025-09-09 07:53:07","extension":"png","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":3478272,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cstrong\u003eFossil locality and Permian outcrop of \u003c/strong\u003e\u003cem\u003e\u003cstrong\u003eDengfengfructus\u003c/strong\u003e\u003c/em\u003e\u003cstrong\u003e gen. nov. near Wangyao village, Gaocheng town, Dengfeng city, Henan Province, (113.155925°E, 34.341893°N).\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003ea. \u003c/strong\u003eThe position of the fossil locality (blue star) in China. \u003cstrong\u003eb. \u003c/strong\u003eThe position of the fossil locality (blue star) in North China. \u003cstrong\u003ec. \u003c/strong\u003eThe position of the fossil locality (blue star) in the suburb of Dengfeng city, Henan Province, China­. \u003cstrong\u003ed. \u003c/strong\u003eFossiliferous strata (the Lower Shihezi (formerly Shihhotse)Formation) dated between 294.4 and 295.1 Ma\u003csup\u003e23\u003c/sup\u003e are sandwiched between the Upper Shihhotse (formerly Upper Shihezi) Formation and the Shanxi Formation. All these formations belong to the Lower Permian. \u003cstrong\u003ee. \u003c/strong\u003eThe outcrop of the Lower Shihezi (formerly Shihhotse)Formation.­ \u003cstrong\u003ef. \u003c/strong\u003eFossiliferous stratum (between the red lines) on the outcrop, enlarged from the rectangle in Fig. 1e.\u003c/p\u003e","description":"","filename":"Figure01.png","url":"https://assets-eu.researchsquare.com/files/rs-7244136/v1/020d2fc4b4f5427c5af6cde5.png"},{"id":90868745,"identity":"3c265520-c3b5-4432-810d-31e61311464a","added_by":"auto","created_at":"2025-09-09 07:53:07","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":10791164,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cstrong\u003eDetailed views of \u003c/strong\u003e\u003cem\u003e\u003cstrong\u003eDengfengfructus maxima \u003c/strong\u003e\u003c/em\u003e\u003cstrong\u003egen. et sp. nov. Holotype. Specimen number PB207444.\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003ea.\u003c/strong\u003e General view of the fruit. Scale bar = 5 mm. \u003cstrong\u003eb. \u003c/strong\u003eTip part of the fruit, enlarged from the top rectangle in Fig. 2a. Note the closed fruit tip. Scale bar = 1 mm.\u003cstrong\u003e c.\u003c/strong\u003e Base part of the fruit, enlarged from the bottom rectangle in Fig. 2a. Note the vascular bundle (black arrow). Scale bar =0.5 mm.\u003cstrong\u003e d.\u003c/strong\u003e Detailed view of the seed/ovule in the fruit, enlarged from the left middle white rectangle in Fig. 2a. Note layered structure (white arrows) of the seed coat and seed content (lower right). Scale bar = 0.5 mm.\u003cstrong\u003e e.\u003c/strong\u003e Detailed view of the seed coat in the fruit, enlarged from the right middle white rectangle in Fig. 2a. Note cellular details of the layered seed coat (black arrows). Scale bar = 0.5 mm.\u003cstrong\u003e f.\u003c/strong\u003e Angular-outlined cells in the seed content, enlarged from the black rectangle in Fig. 2a. Scale bar = 0.5 mm.\u003c/p\u003e","description":"","filename":"Figure02.png","url":"https://assets-eu.researchsquare.com/files/rs-7244136/v1/314167a797e7f5b03a72937f.png"},{"id":90868744,"identity":"e2e39956-8834-4fac-ac49-20620505674e","added_by":"auto","created_at":"2025-09-09 07:53:07","extension":"png","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":9574767,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cstrong\u003eSEM images of \u003c/strong\u003e\u003cem\u003e\u003cstrong\u003eDengfengfructus maxima \u003c/strong\u003e\u003c/em\u003e\u003cstrong\u003egen. et sp. nov. Holotype. Specimen number PB207444.\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003ea.\u003c/strong\u003e Cellular details of the fruit tip, approximately corresponding to the rectangle in Fig. 2b. Scale bar = 0.2 mm. \u003cstrong\u003eb. \u003c/strong\u003eThree layers (1, 2, 3) of the seed coat. Note the borders (arrows) of the layers. Scale bar = 1 mm.\u003cstrong\u003e c.\u003c/strong\u003eThree layers (1, 2, 3) of the seed coat and the seed content (4). Note the borders (arrows) of the layers. Scale bar = 1 mm.\u003cstrong\u003ed.\u003c/strong\u003e Detailed view of the three layers (1, 2, 3) in the rectangle in Fig. 3c. Note the borders (arrows) of the layers. Scale bar = 0.2 mm.\u003c/p\u003e","description":"","filename":"Figure03.png","url":"https://assets-eu.researchsquare.com/files/rs-7244136/v1/89f5b4ccece17abbdab9eac6.png"},{"id":90868907,"identity":"8cf54510-7667-4db5-ae3e-895bef0638c7","added_by":"auto","created_at":"2025-09-09 07:53:24","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":27880654,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-7244136/v1/cd7d15c1-58fd-423e-b013-ab82105b6458.pdf"},{"id":90868747,"identity":"d716b7e2-71a5-4d5b-9b01-71c9c072deaa","added_by":"auto","created_at":"2025-09-09 07:53:07","extension":"tif","order_by":1,"title":"","display":"","copyAsset":false,"role":"supplement","size":9022424,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cstrong\u003eFigure S1 \u003c/strong\u003e\u003cem\u003e\u003cstrong\u003eDengfengfructus maxima \u003c/strong\u003e\u003c/em\u003e\u003cstrong\u003egen. et sp. nov. and associated leaves of \u003c/strong\u003e\u003cem\u003e\u003cstrong\u003eLobatannularia spatulata\u003c/strong\u003e\u003c/em\u003e\u003cstrong\u003e (He) Yang 2006.\u003c/strong\u003e\u003c/p\u003e","description":"","filename":"FigureS1.tif","url":"https://assets-eu.researchsquare.com/files/rs-7244136/v1/899bfe4a9aff27ae74212298.tif"}],"financialInterests":"No competing interests reported.","formattedTitle":"Early Permian fossil fruit reshapes angiosperm history","fulltext":[{"header":"Main text","content":"\u003cp\u003eAlthough angiosperms account for most of the species diversity of plants on Earth\u003csup\u003e1\u003c/sup\u003e, little is known about the origin and early history of angiosperms. The publishing of \u003cem\u003eTaiyuanostachya\u003c/em\u003e\u003csup\u003e16\u003c/sup\u003e and \u003cem\u003eYuzhoua\u003c/em\u003e\u003csup\u003e15\u003c/sup\u003e as Early Permian angiosperms opened a new era of angiosperm study. However, there is a confrontation between some palaeobotanists who insisted on the Early Cretaceous origin for angiosperms\u003csup\u003e10,11\u003c/sup\u003e and the phylogenomic scientists who found a pre-Cretaceous origin for angiosperms\u003csup\u003e7-9\u003c/sup\u003e as well as palaeobotanists who unveiled pre-Cretaceous angiosperms\u003csup\u003e12-17\u003c/sup\u003e. The only reliable way to elucidate the early history of angiosperms is to study more fossils of plants.\u003c/p\u003e\n\u003cp\u003eThe Yuzhou Flora is typical of the Lower Shihezi (formerly Shihhotse)Formation, Lower Permian in Henan, China\u003csup\u003e18\u003c/sup\u003e, and is one of the best studied parts of the Cathayasian Flora\u003csup\u003e19-21\u003c/sup\u003e. Previous studies indicate that the Yuzhou Flora comprises 307 species in 111 genera of Sphenophyllales, Equisetales, Filicales, Lepidodendrales, Noeggerathiales, Peltaspermales, Pteridospermophytes, Cycadales, Dicranohpyllopsida, Ginkgophyta, Cordaitopsida, Coniferopsida, Gigantopteridales, and plants incertae sedis\u003csup\u003e18\u003c/sup\u003e. The age of this flora has been dated as the Kungurian–Roadian\u0026nbsp;Asselian\u003csup\u003e22\u003c/sup\u003e, namely, 294.4–295.1 Ma\u003csup\u003e23\u003c/sup\u003e.\u0026nbsp;The specimen was collected from an outcrop of the Lower Shihezi (formerly Shihhotse)Formation near Wangyao\u0026nbsp;village, Gaocheng\u0026nbsp;town, Dengfeng\u0026nbsp;city, Henan Province, China (113.155925°E, 34.341893°N; Fig. 1) by Y.L. in May, 2025. The fossil was exposed on the surface naturally, requiring little trimming or dégagement. The morphology was photographed using a Nikon D800. The details of the specimen were photographed\u0026nbsp;via\u0026nbsp;a Zeiss Zoom V16 stereomicroscope.\u0026nbsp;All the figures were organized for publication in Photoshop 7.0 software.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eAngiosperms\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eOrder \u003cem\u003eincertae sedis\u003c/em\u003e\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eFamily \u003cem\u003eincertae sedis\u003c/em\u003e\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eDengfengfructus\u0026nbsp;\u003c/em\u003egen. nov.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eType species\u003c/strong\u003e: \u003cem\u003eDengfengfructus maxima\u003c/em\u003e gen. et sp. nov.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eDiagnosis\u003c/strong\u003e: Drupe oval, enclosing a single seed. Seed comprising a three-layered seed coat and seed content.\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eDengfengfructus maxima\u003c/em\u003e gen. et sp. nov.\u003c/p\u003e\n\u003cp\u003e(Figures 2–3)\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eDiagnosis\u003c/strong\u003e: The same as that of the genus, for the time being.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eDescription:\u0026nbsp;\u003c/strong\u003eThe fruit is preserved as a compression, oval–shaped, 17.6 mm long and 14 mm\u0026nbsp;wide (Fig. 2a). The fruit comprises a seed and a surrounding pericarp (Fig. 2a). The thickness of the pericarp varies from 2 to 3.3 mm (Fig. 2a). The pericarp appears homogeneous in structure, with a basal vascular bundle and secluded tip, with cellular details (Figs. 2a-c, 3a). The seed is approximately 10 mm in diameter and 12 mm tall, oval–shaped, and surrounded by a three-layered seed coat containing a seed content that is in turn fully enclosed by the pericarp (Figs. 2a, d-e, 3b-d). The three layers of the seed coat are 470 μm, 390 μm, and 240 μm\u0026nbsp;thick,\u0026nbsp;respectively, from inner to outer (Fig. 2e). The cells in the seed coat have conspicuous angular outlines, 82~210\u0026nbsp;μm\u0026nbsp;×\u0026nbsp;145~424 μm, with little differentiation among the layers (Fig. 2e). In contrast, the seed content is homogeneous, including cells with inconspicuous outlines\u0026nbsp;of\u0026nbsp;75~118 μm\u0026nbsp;×\u0026nbsp;101~193 μm (Fig. 2f). The fruit is associated with\u0026nbsp;the\u0026nbsp;leaves of \u003cem\u003eLobatannularia spatulata\u0026nbsp;\u003c/em\u003e(He) Yang (Fig. S1a).\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eEtymology\u003c/strong\u003e: \u003cem\u003eDengfengfructus\u003c/em\u003e is for a fruit from the city of Dengfeng in Henan, China, where the fossil is from. The specific epithet \u003cem\u003emaxima\u003c/em\u003e is for the large size of the fruit.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eHolotype specimen:\u0026nbsp;\u003c/strong\u003ePB207444.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eDepository:\u0026nbsp;\u003c/strong\u003eNanjing Institute of Geology and Palaeontology, Nanjing, China.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eType locality\u003c/strong\u003e: Wangyao village, Gaocheng town, Dengfeng city, Henan Province, China (113.155925°E, 34.341893°N).\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eHorizon\u003c/strong\u003e: the Lower Shihezi (formerly Shihhotse)Formation, Permian (294.4-295.1 Ma)\u003csup\u003e23\u003c/sup\u003e.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eRemarks:\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe age of our fossil is supported by its associated leaves, \u003cem\u003eLobatannularia spatulata\u0026nbsp;\u003c/em\u003e(He) Yang 2006, preserved on the same specimen (Fig. S1a). \u003cem\u003eLobatannularia spatulata\u0026nbsp;\u003c/em\u003e(He) Yang is frequently seen in the Yuzhou Flora and has been well studied in previous works\u003csup\u003e18\u003c/sup\u003e. The co-occurrence of these two fossils on the same piece of specimen, on one hand, is suggestive of the abundant occurrence of plant fossils in the strata; on the other hand, the repeated occurrences of \u003cem\u003eLobatannularia spatulata\u0026nbsp;\u003c/em\u003e(He) Yang in various studies pin down the Early Permian age for \u003cem\u003eDengfengfructus\u003c/em\u003e\u003csup\u003e18\u003c/sup\u003e. Two recent thorough stratigraphic surveys\u003csup\u003e22,23\u003c/sup\u003e suggested a Kungurian–Roadian\u0026nbsp;Asselian age (294.4-295.1 Ma) for our fossil plant.\u003c/p\u003e\n\u003cp\u003eThe single seed of \u003cem\u003eDengfengfructus\u003c/em\u003e is fully enclosed by a pericarp, satisfying the original criterion for angiosperms given by Paul Hermann in 1690\u003csup\u003e24\u003c/sup\u003e. This is the main reason we interpret \u003cem\u003eDengfengfructus\u0026nbsp;\u003c/em\u003eas an angiosperm fruit. However, considering Palaeozoic angiosperms are still rare and that seeds are most likely present in the Early Permian, we prefer to exclude all alternative interpretations before proceeding further.\u003c/p\u003e\n\u003cp\u003eTheoval general morphology of \u003cem\u003eDengfengfructus\u003c/em\u003e distinguishes itself from all known vegetative parts of plants. Among the reproductive organs of plants, there are no male organs of similar size or organization. Therefore, we do not consider any male reproductive organs in the following discussion. The most plausible alternatives for\u0026nbsp;\u003cem\u003eDengfengfructus\u003c/em\u003e are a female reproductive organ or its part (namely, a fruit, an ovule, or a seed).\u003c/p\u003e\n\u003cp\u003eSince ovules and seeds were observed as early as the Late Devonian, we must consider these alternatives seriously for our Permian fossil organ. The morphology and organization of Palaeozoic ovules/seeds are well known\u003csup\u003e25\u003c/sup\u003e. A mature Palaeozoic seed usually has a seed content surrounded by a three-layered seed coat (including sarcotesta, sclerotesta, and endotesta)\u003csup\u003e25\u003c/sup\u003e. In contrast to seed content, which is unlikely to be preserved in fossils, a seed coat is the part that has the greatest potential to be fossilized in a seed. This generalization contrasts with our observations: the peripheral surrounding layer of \u003cem\u003eDengfengfructus\u003c/em\u003e is homogeneous and shows no trace of stratification (Fig. 2a); thus, it is distinct from any known seed coat. Apparently, \u003cem\u003eDengfengfructus\u003c/em\u003e cannot be compared to any mature seeds of any plant. How about an immature seed? An immature seed usually shows a smooth and gradual transition to an ovule in morphology and organization. An ovule usually includes a nucellus and protective integuments. Integuments (corresponding to the seed coat in a mature seed) partially surround the nucellus and may appear homogeneous when immature or may demonstrate stratification when mature, whereas the nucellus corresponds to the seed content in a mature seed and usually demonstrates no stratification. This again contradicts our observation: the central oval body in \u003cem\u003eDengfengfructus\u003c/em\u003e apparently has a three-layered peripheral part and a central homogeneous content (Figs. 2a, d-f, 3b-d). By the way, the seeds of interesting plants in Bennetitales have their characteristic micropylar tubes, excluding themselves from our consideration. The above contrasts and comparisons exclude the possibility of \u003cem\u003eDengfengfructus\u003c/em\u003e being either a seed or an ovule, leaving only one alternative for \u003cem\u003eDengfengfructus\u003c/em\u003e: fruit.\u003c/p\u003e\n\u003cp\u003eAlthough a Permian angiosperm is unacceptable for many, as it is far older than the widely accepted age for angiosperms (the Early Cretaceous\u003csup\u003e10,11\u003c/sup\u003e), recent phylogenomic studies and palaeobotanical studies of \u003cem\u003eTaiyuanostachya\u003c/em\u003e\u003csup\u003e16\u003c/sup\u003e and \u003cem\u003eYuzhoua\u003c/em\u003e\u003csup\u003e15\u003c/sup\u003e have undermined the credibility of the Cretaceous origin for angiosperms\u003csup\u003e10,11\u003c/sup\u003e, making the angiospermous affinity of \u003cem\u003eDengfengfructus\u0026nbsp;\u003c/em\u003eless surprising.\u003c/p\u003e\n\u003cp\u003eAccording to the traditional thought dating back to the age of Arber and Parkin (1907), ancestral carpels are conduplicate carpels with marginal ovules\u003csup\u003e26\u003c/sup\u003e. However, this expectation is not honored by fossil evidence. For example, although Dilcher and Crane\u003csup\u003e27\u003c/sup\u003e appeared to support traditional thinking via their fossil evidence of \u003cem\u003eArachaeanthus\u003c/em\u003e, subsequent careful examination indicated that the seeds/ovules in \u003cem\u003eArchaeanthus\u003c/em\u003e are actually borne at the dorsal margin of the fruit/carpel, thus refuting the traditional thinking\u003csup\u003e28\u003c/sup\u003e. In our case, the basally fixed seed in \u003cem\u003eDengfengfructus\u003c/em\u003e is apparently at odds with the widely expected marginally-positioned seeds, further reducing the credibility of the traditional thinking and lending support to the recently advanced Unifying Theory\u003csup\u003e29,30\u003c/sup\u003e, which allowed various positions of a seed in a fruit. Notably, even studies on carpels of the assumed ancestral living Magnoliales (\u003cem\u003eMichelia\u003c/em\u003e\u003csup\u003e31\u003c/sup\u003e and \u003cem\u003eIllicium\u003c/em\u003e\u003csup\u003e32\u003c/sup\u003e) have rejected the traditional thinking. On the basis of currently available data, the traditional thinking of angiosperm evolution, although widely accepted and taught in classrooms, is highly flawed and should be discarded.\u003c/p\u003e\n\u003cp\u003eA caveat of the current conclusion that \u003cem\u003eDengfengfructus\u003c/em\u003e is an angiosperm, the pollination process of which is unknown. Since Tomlinson and Takaso\u003csup\u003e33\u003c/sup\u003e have distinguished angiosperms from gymnosperms by the ovule-enclosing status of a plant at the time of pollination, namely, ovule-enclosing occurs before pollination in angiosperms. This reduces the validity of our above judgement, but it does not reduce the evolutionary significance of \u003cem\u003eDengfengfructus.\u0026nbsp;\u003c/em\u003eThere are only two possible scenarios in the future. 1) If a future study proves that ovule enclosure occurs before pollination in \u003cem\u003eDengfengfructus\u003c/em\u003e, then \u003cem\u003eDengfengfructus\u003c/em\u003e will be confirmed as a \u003cem\u003ebona fide\u003c/em\u003e Palaeozoic angiosperm. In this case, the existence of angiosperms in the Early Permian will be verified and consolidated, and angiosperm diversity in the Permian (Palaeozoic) is further underscored, especially when \u003cem\u003eTaiyuanostachya\u003c/em\u003e\u003csup\u003e16\u003c/sup\u003e and \u003cem\u003eYuzhoua\u003c/em\u003e\u003csup\u003e15\u003c/sup\u003e are taken into consideration. 2) However, if a future study proves that ovule/seed\u0026nbsp;enclosure\u0026nbsp;in \u003cem\u003eDengfengfructus\u003c/em\u003e occurs after pollination, then \u003cem\u003eDengfengfructus\u003c/em\u003e should be placed among gymnosperms. If \u003cem\u003eDengfengfructus\u003c/em\u003e were kicked out of angiosperms, it would not affect the Permian origin of angiosperms: \u003cem\u003eTaiyuanostachya\u003c/em\u003e has not only demonstrates not only angiospermy (enclosed seed) but also angio-ovuly (enclosed ovule), thus having pushed the origin time of angiosperms into the EarlyPermian\u003csup\u003e16\u003c/sup\u003e. This alternative placement of \u003cem\u003eDengfengfructus\u0026nbsp;\u003c/em\u003emeans that \u003cem\u003eDengfengfructus\u003c/em\u003e, although a gymnosperm, has developed a seed-bearing structure that is hard to\u0026nbsp;distinguish\u0026nbsp;from an angiosperm fruit. This would reduce the former huge gap between angiosperms and gymnosperms, underscore the paper-thin and blurry boundary between angiosperms and gymnosperms, and\u0026nbsp;corroborate\u0026nbsp;the theoretically predicted smooth transition between angiosperms and gymnosperms.\u003c/p\u003e\n\u003cp\u003eThe timing order of two independent developmental events, ovule-enclosing and pollination, matters in terms of the taxonomy of seed plants and is the key\u0026nbsp;feature distinguishing gymnosperms and angiosperms. During the long-term\u0026nbsp;evolution of seed plants, some of them may arbitrarily sport\u0026nbsp;and\u0026nbsp;develop earlier ovule\u0026nbsp;closure. This initially randomly occurring but later canalized and fixed feature apparently agrees with the general pattern of organism evolution,\u0026nbsp;namely, offspring development conditioning\u0026nbsp;(ODC)\u003csup\u003e34\u003c/sup\u003e. If such a feature benefits the plants in terms of proliferation, it is very likely to be canalized and carried on by their offspring. This developmental pattern may be heritable in plants. The end product of such a process is the debut of angiosperms in geological history.\u003c/p\u003e"},{"header":"Conclusions","content":"\u003cp\u003e\u003cem\u003eDengfengfructus maxima\u003c/em\u003e gen. et sp. nov. is a new Permian angiosperm fruit. Its Early Permian (294.4–295.1 Ma) age confirms the existence of angiosperms in the late Palaeozoic. The margin-independent position of seed in \u003cem\u003eDengfengfructus\u003c/em\u003e implies that the traditional thinking of angiosperms, which has been refuted by previous studies, is not favoured by our latest fossil evidence. The doubts cast over traditional thinking by various studies imply that we are facing a new wave of innovations in botanical theories, especially those concerned with angiosperm origin and evolution.\u003c/p\u003e\n\u003cp\u003e\u003cbr\u003e\u003c/p\u003e"},{"header":"Abbreviations","content":"\u003cp\u003eSEM (scanning electron microscope)\u003c/p\u003e\n\u003cp\u003e\u003cbr\u003e\u003c/p\u003e"},{"header":"Declarations","content":"\u003cp\u003e\u003cstrong\u003eEthics approval and consent to participate\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eNot applicable\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eConsent for publication\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eNot applicable\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eFunding\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe National Key Research and Development Program of China (2024YFF0807601).\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eAuthors\u0026apos; contributions\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eWH and YL collected the samples. XW initiated the study and drafted the manuscript. QF, WH, YL and XW analysed the data and modified and finalized the manuscript.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eAcknowledgements\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eWe appreciate Ms. Jingjing Tang for assisting with the photography, and Mr. Yan Fang for assisting with SEM observation.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eData availability\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eAll the data analysed during this study are included in this article.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eCompeting interests\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe authors declare that they have no competing interests.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\u003cli\u003e\u003cspan\u003eCrepet WL, Niklas KJ. Darwin's second abominable mystery: Why are there so many angiosperm species? 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J Northwest Univ (Natural Sci Edition). 2021;51:163\u0026ndash;72. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.16152/j.cnki.xdxbzr.2021-01-018\u003c/span\u003e\u003cspan address=\"10.16152/j.cnki.xdxbzr.2021-01-018\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e.\u003c/span\u003e\u003c/li\u003e\u003c/ol\u003e"},{"header":"Tables","content":"\u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab1\" border=\"1\"\u003e\u003cdiv category=\"Completeness\" id=\"6\" ruleid=\"MissingTableCitation_01\" status=\"ignored\" values=\"Table 1\" class=\"btn-xs-small Annotation tooltipped\" data-position=\"top\" data-tooltip=\"\"\u003e\u003c/div\u003e\u003ccaption language=\"En\"\u003e\u003cdiv class=\"CaptionNumber\"\u003eTable 1\u003c/div\u003e\u003cdiv class=\"CaptionContent\"\u003e\u003cdiv class=\"SimplePara\"\u003eComparison among contemporaneous \u003cspan type=\"Italic\" class=\"Italic\" name=\"Emphasis\"\u003eDengfengfructus, Protofructus, Novofructus, Taiyuanostachya\u003c/span\u003e, and \u003cspan type=\"Italic\" class=\"Italic\" name=\"Emphasis\"\u003eYuzhoua\u003c/span\u003e.\u003c/div\u003e\u003c/div\u003e\u003c/caption\u003e\u003ccolgroup cols=\"6\"\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e\u003cthead\u003e\u003ctr\u003e\u003cth align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/th\u003e\u003cth align=\"left\" colname=\"c2\"\u003e\u003cdiv class=\"SimplePara\"\u003e\u003cspan type=\"Italic\" class=\"Italic\" name=\"Emphasis\"\u003eDengfengfructus\u003c/span\u003e\u003c/div\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c3\"\u003e\u003cdiv class=\"SimplePara\"\u003e\u003cspan type=\"Italic\" class=\"Italic\" name=\"Emphasis\"\u003eProtofructus\u003c/span\u003e\u003c/div\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c4\"\u003e\u003cdiv class=\"SimplePara\"\u003e\u003cspan type=\"Italic\" class=\"Italic\" name=\"Emphasis\"\u003eNovofructus\u003c/span\u003e\u003c/div\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c5\"\u003e\u003cdiv class=\"SimplePara\"\u003e\u003cspan type=\"Italic\" class=\"Italic\" name=\"Emphasis\"\u003eTaiyuanostachya\u003c/span\u003e\u003c/div\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c6\"\u003e\u003cdiv class=\"SimplePara\"\u003e\u003cspan type=\"Italic\" class=\"Italic\" name=\"Emphasis\"\u003eYuzhoua\u003c/span\u003e\u003c/div\u003e\u003c/th\u003e\u003c/tr\u003e\u003c/thead\u003e\u003ctbody\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cdiv class=\"SimplePara\"\u003ePreservation\u003c/div\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cdiv class=\"SimplePara\"\u003eCompression\u003c/div\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cdiv class=\"SimplePara\"\u003eCompression\u003c/div\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cdiv class=\"SimplePara\"\u003eCompression\u003c/div\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cdiv class=\"SimplePara\"\u003eCompression\u003c/div\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cdiv class=\"SimplePara\"\u003eCompression\u003c/div\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cdiv class=\"SimplePara\"\u003eFossil locality\u003c/div\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cdiv class=\"SimplePara\"\u003eDengfeng, Henan\u003c/div\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cdiv class=\"SimplePara\"\u003eXinmi, Henan\u003c/div\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cdiv class=\"SimplePara\"\u003eXinmi, Henan\u003c/div\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cdiv class=\"SimplePara\"\u003eTaiyuan, Shanxi\u003c/div\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cdiv class=\"SimplePara\"\u003eYuzhou, Henan\u003c/div\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cdiv class=\"SimplePara\"\u003eAge\u003c/div\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cdiv class=\"SimplePara\"\u003eEarly Permian\u003c/div\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cdiv class=\"SimplePara\"\u003eEarly Permian\u003c/div\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cdiv class=\"SimplePara\"\u003eEarly Permian\u003c/div\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cdiv class=\"SimplePara\"\u003eEarly Permian\u003c/div\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cdiv class=\"SimplePara\"\u003eEarly Permian\u003c/div\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cdiv class=\"SimplePara\"\u003eHorizon\u003c/div\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"5\" nameend=\"c6\" namest=\"c2\"\u003e\u003cdiv class=\"SimplePara\"\u003eLower Shihezi (formerly Shihhotse)Fm.\u003c/div\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cdiv class=\"SimplePara\"\u003eShape of organ\u003c/div\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cdiv class=\"SimplePara\"\u003eoval\u003c/div\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cdiv class=\"SimplePara\"\u003eelongated\u003c/div\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cdiv class=\"SimplePara\"\u003eelongated\u003c/div\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cdiv class=\"SimplePara\"\u003ecylindrical\u003c/div\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cdiv class=\"SimplePara\"\u003ebulging\u003c/div\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cdiv class=\"SimplePara\"\u003eOrganization of organ\u003c/div\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cdiv class=\"SimplePara\"\u003esole fruit\u003c/div\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cdiv class=\"SimplePara\"\u003esole fruit\u003c/div\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cdiv class=\"SimplePara\"\u003esole fruit\u003c/div\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cdiv class=\"SimplePara\"\u003einfructescence\u003c/div\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cdiv class=\"SimplePara\"\u003esole fruit\u003c/div\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cdiv class=\"SimplePara\"\u003eLength of organ\u003c/div\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cdiv class=\"SimplePara\"\u003e17.6 mm\u003c/div\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cdiv class=\"SimplePara\"\u003e43 mm\u003c/div\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cdiv class=\"SimplePara\"\u003e36 mm\u003c/div\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cdiv class=\"SimplePara\"\u003e43 mm\u003c/div\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cdiv class=\"SimplePara\"\u003e40 mm\u003c/div\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cdiv class=\"SimplePara\"\u003eOvules per fruit\u003c/div\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cdiv class=\"SimplePara\"\u003e1\u003c/div\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cdiv class=\"SimplePara\"\u003e8+\u003c/div\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cdiv class=\"SimplePara\"\u003e2+\u003c/div\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cdiv class=\"SimplePara\"\u003e1\u003c/div\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cdiv class=\"SimplePara\"\u003e3\u003c/div\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cdiv class=\"SimplePara\"\u003eOvule position\u003c/div\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cdiv class=\"SimplePara\"\u003ebasal\u003c/div\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cdiv class=\"SimplePara\"\u003eon a long placenta\u003c/div\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cdiv class=\"SimplePara\"\u003ealong fruit margin\u003c/div\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cdiv class=\"SimplePara\"\u003ebasal in fruit,\u003c/div\u003e\u003cdiv class=\"SimplePara\"\u003eon infructescence axis\u003c/div\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cdiv class=\"SimplePara\"\u003eon side ovarian wall\u003c/div\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cdiv class=\"SimplePara\"\u003eNumber of specimens\u003c/div\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cdiv class=\"SimplePara\"\u003e1\u003c/div\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cdiv class=\"SimplePara\"\u003e1\u003c/div\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cdiv class=\"SimplePara\"\u003e1\u003c/div\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cdiv class=\"SimplePara\"\u003e6\u003c/div\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cdiv class=\"SimplePara\"\u003e6\u003c/div\u003e\u003c/td\u003e\u003c/tr\u003e\u003c/tbody\u003e\u003c/colgroup\u003e\u003c/table\u003e\u003c/div\u003e\u003cbr/\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":true,"hideJournal":true,"highlight":"","institution":"","isAcceptedByJournal":false,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"researchsquare","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":true,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"/submission","title":"Research Square","twitterHandle":"researchsquare","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"","reportingPortfolio":"","inReviewEnabled":false,"inReviewRevisionsEnabled":true},"keywords":"angiosperm, fruit, Permian, Palaeozoic, 294.4 Ma","lastPublishedDoi":"10.21203/rs.3.rs-7244136/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-7244136/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eAngiosperms are the most important plant group for humans in the current earth ecosystem\u003csup\u003e\u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e1\u003c/span\u003e\u003c/sup\u003e. However, the origin and early history of angiosperms remain elusive and controversial \u003csup\u003e\u003cspan additionalcitationids=\"CR3 CR4 CR5 CR6 CR7 CR8\" citationid=\"CR2\" class=\"CitationRef\"\u003e2\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e9\u003c/span\u003e\u003c/sup\u003e. According to the over60-year-old stereotype in botany, angiosperms cannot be older than the Cretaceous\u003csup\u003e\u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e10\u003c/span\u003e,\u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e11\u003c/span\u003e\u003c/sup\u003e. However, this thinking is now facing increasing challenges in terms of fossil evidence\u003csup\u003e\u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e6\u003c/span\u003e,\u003cspan additionalcitationids=\"CR13 CR14 CR15 CR16\" citationid=\"CR12\" class=\"CitationRef\"\u003e12\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e17\u003c/span\u003e\u003c/sup\u003e and molecular estimates\u003csup\u003e\u003cspan additionalcitationids=\"CR8\" citationid=\"CR7\" class=\"CitationRef\"\u003e7\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e9\u003c/span\u003e\u003c/sup\u003e. Apparently, serious and independent studies on new early fossil angiosperms are urgently needed to resolve this academic confrontation. In recent years, a renewed wave of interest in Permian fossil plants in Henan\u0026rsquo;s Cathaysian flora has been rekindled by the latest palaeobotanical progress\u003csup\u003e\u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e15\u003c/span\u003e,\u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e16\u003c/span\u003e\u003c/sup\u003e. During a recent field excursion in May 2025, we collected a new fossil organ from the Lower Shihezi (formerly Shihhotse)Formation (Lower Permian) of Dengfeng, Henan, China. This new fossil organ is interpreted as a large fruit and named \u003cem\u003eDengfengfructus maxima\u003c/em\u003e gen. et sp. nov. The seed enclosed in the fruit has a peripheral three-layered seed coat, which distinguishes the seed itself from an otherwise possible nucellus in a seed. The good preservation allows the cellular details in the seed coat and seed content to be revealed. Unlike contemporary \u003cem\u003eTaiyuanostachya\u003c/em\u003e\u003csup\u003e\u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e16\u003c/span\u003e\u003c/sup\u003e and \u003cem\u003eYuzhoua\u003c/em\u003e\u003csup\u003e\u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e15\u003c/span\u003e\u003c/sup\u003e, \u003cem\u003eDengfengfructus\u003c/em\u003e has a large size and a thick pericarp. This new fossil organ apparently updates and enhances the current understanding of angiosperms and their diversity in the Permian. The history of angiosperms can be dated back to the Early Permian (Palaeozoic). The former stereotype that angiosperms are restricted to the Cretaceous and later ages falls apart in front of the currently available palaeobotanical progress and molecular estimates. Obviously, we are facing a wave of botanical theories in the near future.\u003c/p\u003e","manuscriptTitle":"Early Permian fossil fruit reshapes angiosperm history","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2025-09-09 07:53:02","doi":"10.21203/rs.3.rs-7244136/v1","editorialEvents":[{"type":"communityComments","content":0}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"researchsquare","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":true,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"/submission","title":"Research Square","twitterHandle":"researchsquare","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"","reportingPortfolio":"","inReviewEnabled":false,"inReviewRevisionsEnabled":true}}],"origin":"","ownerIdentity":"bdd64f22-ed3b-42b1-80b5-d039a4f63fd8","owner":[],"postedDate":"September 9th, 2025","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"posted","subjectAreas":[],"tags":[],"updatedAt":"2025-09-09T07:53:02+00:00","versionOfRecord":[],"versionCreatedAt":"2025-09-09 07:53:02","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-7244136","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-7244136","identity":"rs-7244136","version":["v1"]},"buildId":"8U1c8b4HqxoKbykW_rLl7","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}