{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,6,18]],"date-time":"2026-06-18T09:13:39Z","timestamp":1781774019759,"version":"3.54.5"},"reference-count":53,"publisher":"Oxford University Press (OUP)","issue":"6","license":[{"start":{"date-parts":[[2024,11,9]],"date-time":"2024-11-09T00:00:00Z","timestamp":1731110400000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/academic.oup.com\/pages\/standard-publication-reuse-rights"}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"published-print":{"date-parts":[[2026,6,17]]},"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:sec>\n                    <jats:title>Background and Aims<\/jats:title>\n                    <jats:p>Eocene foliage formerly attributed to the extant araliaceous genus Oreopanax was found attached to twigs bearing inflorescences and infructescences unlike those of Araliaceae. Using newly observable characters of phyllotaxy, vegetative and floral buds, infructescences and seeds, we sought to reassess the affinities of this strange angiosperm.<\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Methods<\/jats:title>\n                    <jats:p>Fossils were studied from the Parachute Creek Member of the Green River Formation from near Bonanza, Utah and Douglas Pass, Colorado (~47 Mya). Macrofossil impression remains were investigated by low-angle reflected light, and subtle details of the vegetative and floral buds, stamens, mature fruits and seeds were revealed by optical shadow effect microscopy, documenting previously obscure topographic surface features.<\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Key Results<\/jats:title>\n                    <jats:p>Othniophyton elongatum (MacGinitie) Manchester, Judd, Correa-Narvaez gen. et comb. nov. has simple, short-petiolate, elongate, entire-margined leaves with thick midveins, pinnate, brochidodromous secondaries, common intersecondary veins and finely reticulate higher-order venation. Inflorescences are small axillary cymes; flower buds are subglobose and pedicellate, in bract axils. The flowers are actinomorphic, bisexual, with approximately five imbricate perianth parts, ~24 stamens with elongate anthers and short filaments, arising from a hypanthium. The whorl of stamens persists to fruiting stage. The ovary is superior, with probable basal placentation and five stigmatic arms. Fruits are pedicellate berries with a cup-shaped hypanthium and contain ~15 lensoidal reniform seeds, each with a curved embryo and ornamented with concentric ridges. The combined characters refute the prior placement in Araliaceae and rule out affinities with most extant clades of angiosperms.<\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Conclusions<\/jats:title>\n                    <jats:p>The distinctive combination of observed features does not coincide with any extant family. Among extant orders of Eudicots, the fossil seems to conform most closely to the order Caryophyllales, but key differences remain. This example indicates that the vegetation of ~47 Mya included some taxa that cannot readily be placed in modern families and genera.<\/jats:p>\n                  <\/jats:sec>","DOI":"10.1093\/aob\/mcae196","type":"journal-article","created":{"date-parts":[[2024,11,7]],"date-time":"2024-11-07T15:26:50Z","timestamp":1730993210000},"page":"1971-1986","source":"Crossref","is-referenced-by-count":3,"title":["Vegetative and reproductive morphology of\n                    <i>Othniophyton elongatum<\/i>\n                    (MacGinitie) gen. et comb. nov., an extinct angiosperm of possible caryophyllalean affinity from the Eocene of Colorado and Utah, USA"],"prefix":"10.1093","volume":"137","author":[{"given":"Steven R","family":"Manchester","sequence":"first","affiliation":[{"name":"Florida Museum of Natural History, Dickinson Hall , Gainesville, FL 32611-7800 ,","place":["USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Walter S","family":"Judd","sequence":"additional","affiliation":[{"name":"Bell Museum Herbarium, University of Minnesota , Saint Paul, MN 55108-0215 ,","place":["USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Julian E","family":"Correa-Narvaez","sequence":"additional","affiliation":[{"name":"Florida Museum of Natural History, Dickinson Hall , Gainesville, FL 32611-7800 ,","place":["USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"286","published-online":{"date-parts":[[2024,11,9]]},"reference":[{"key":"2026061716492711000_CIT0001","first-page":"1","article-title":"An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG IV","volume":"181","author":"APG IV","year":"2016","journal-title":"Botanical Journal of the Linnean Society"},{"key":"2026061716492711000_CIT0002","doi-asserted-by":"crossref","first-page":"5","DOI":"10.3100\/hpib.v21iss1.2016.n2","article-title":"First report of Rhabdodendraceae for the vascular flora of Colombia and the Upper R\u00edo Negro basin, with comments on phytogeography, habitats, and distribution of Rhabdodendron amazonicum","volume":"21","author":"Aymard","year":"2016","journal-title":"Harvard Papers in Botany"},{"key":"2026061716492711000_CIT0042","doi-asserted-by":"crossref","first-page":"635","DOI":"10.1111\/j.1654-1103.2004.tb02305.x","article-title":"Relationships between plant traits and climate in the Mediterranean region: a pollen data analysis","volume":"15","author":"Barboni","year":"2004","journal-title":"Journal of Vegetation Science"},{"key":"2026061716492711000_CIT0044","first-page":"55","volume":"165-B","author":"Berry","year":"1930","journal-title":"A flora of Green River age in the Wind River Basin of Wyoming"},{"key":"2026061716492711000_CIT0003","doi-asserted-by":"crossref","first-page":"1389","DOI":"10.3732\/ajb.90.9.1389","article-title":"An extinct genus of Salicaceae based on twigs with attached flowers, fruits, and foliage from the Eocene Green River Formation of Utah and Colorado, USA","volume":"90","author":"Boucher","year":"2003","journal-title":"American Journal of Botany"},{"key":"2026061716492711000_CIT0004","doi-asserted-by":"crossref","first-page":"496","DOI":"10.1080\/10635150490445913","article-title":"Molecular phylogenetic dating of asterid flowering plants shows early Cretaceous diversification","volume":"53","author":"Bremer","year":"2004","journal-title":"Systematic Biology"},{"key":"2026061716492711000_CIT0005","first-page":"279","article-title":"Additions to the flora of the Green River Formation","volume":"154-J","author":"Brown","year":"1929","journal-title":"U.S. Geological Survey Professional Paper"},{"key":"2026061716492711000_CIT0006","first-page":"44","article-title":"The recognizable species of the Green River flora","volume":"185-C","author":"Brown","year":"1934","journal-title":"U.S. Geological Survey Professional Paper"},{"key":"2026061716492711000_CIT0007","doi-asserted-by":"crossref","first-page":"305","DOI":"10.1016\/0034-6667(94)90007-8","article-title":"Parvileguminophyllum coloradensis, a new combination for Mimosites coloradensis Knowlton, Green River Formation of Utah and Colorado","volume":"80","author":"Call","year":"1994","journal-title":"Review of Palaeobotany and Palynology"},{"key":"2026061716492711000_CIT0009","doi-asserted-by":"crossref","DOI":"10.7208\/chicago\/9780226536705.001.0001","volume-title":"Plants of the world. An illustrated encyclopedia of vascular plants","author":"Christenhusz","year":"2017"},{"key":"2026061716492711000_CIT0045","doi-asserted-by":"crossref","first-page":"1","DOI":"10.5479\/si.00963801.66-2556.1","article-title":"Plant and insect fossils from the Green River Eocene of Colorado","volume":"66","author":"Cockerell","year":"1925","journal-title":"Proceedings of the United States National Museum"},{"key":"2026061716492711000_CIT0010","volume-title":"An integrated system of classification of flowering plants","author":"Cronquist","year":"1981"},{"key":"2026061716492711000_CIT0011","author":"Dallwitz","year":"1993","journal-title":"User\u2019s guide to the DELTA System: a general system for processing taxonomic descriptions"},{"key":"2026061716492711000_CIT0012","doi-asserted-by":"publisher","first-page":"645","DOI":"10.5852\/ejt.2020.645","article-title":"Survey of the fruits and endocarps of Icacinaceae (Lamiids, Icacinales)","author":"Del Rio","year":"2020","journal-title":"European Journal of Taxonomy"},{"key":"2026061716492711000_CIT0013","doi-asserted-by":"crossref","first-page":"dsac051","DOI":"10.1093\/dnares\/dsac051","article-title":"Karyotype evolution of the Asterids insights from the first genome sequences of the family Cornaceae","volume":"30","author":"Dong","year":"2023","journal-title":"DNA Research"},{"key":"2026061716492711000_CIT0014","doi-asserted-by":"crossref","first-page":"923","DOI":"10.2307\/4117819","article-title":"Familial relationships of Rhabdodendron (Rhabdodendraceae): plastid rbcL sequences indicate a caryophyllid placement","volume":"52","author":"Fay","year":"1997","journal-title":"Kew Bulletin"},{"key":"2026061716492711000_CIT0015","doi-asserted-by":"crossref","first-page":"121","DOI":"10.1007\/978-4-431-65918-1_6","article-title":"Fossil history of","volume-title":"Evolution and diversification of land plants","author":"Friis","year":"1997"},{"key":"2026061716492711000_CIT0016","doi-asserted-by":"crossref","DOI":"10.1017\/CBO9780511980206","volume-title":"Early flowers and angiosperm evolution","author":"Friis","year":"2011"},{"key":"2026061716492711000_CIT0017","doi-asserted-by":"crossref","DOI":"10.7208\/chicago\/9780226922980.001.0001","volume-title":"The lost world of Fossil Lake: snapshots from deep time","author":"Grande","year":"2013"},{"key":"2026061716492711000_CIT0018","first-page":"261","volume-title":"Determination of angiosperm families by means of a punched-card system","author":"Hansen","year":"1969"},{"key":"2026061716492711000_CIT0048","doi-asserted-by":"crossref","first-page":"138","DOI":"10.1086\/728411","article-title":"Revisiting the Iconic Macginitiea plant and its implications for biogeography, basilaminar lobe development, and evolution in Platanaceae","volume":"185","author":"Huegele","year":"2024","journal-title":"International Journal of Plant Sciences"},{"key":"2026061716492711000_CIT0020","first-page":"53","article-title":"Vegetational characteristics in Europe around the late early to early middle Miocene based on the plant macro record","volume":"249","author":"Jechorek","year":"2004","journal-title":"Courier Forschungsinstitut Senckenberg"},{"key":"2026061716492711000_CIT0019","doi-asserted-by":"crossref","first-page":"747","DOI":"10.1007\/s10265-015-0739-2","article-title":"First occurrence of Cedrelospermum (Ulmaceae) in Asia and its biogeographic implications","volume":"128","author":"Jia","year":"2015","journal-title":"Journal of Plant Research"},{"key":"2026061716492711000_CIT0021","first-page":"329","article-title":"Megaflora of the Hell Creek and lower Fort Union Formations in the western Dakotas: vegetational response to climate change, the Cretaceous-Tertiary boundary event, and rapid marine transgression. In Hartman JH, Johnson KR, Nichols DJ eds. The Hell Creek Formation and the Cretaceous-Tertiary Boundary in the Northern Great Plains: An integrated continental record of the end of the Cretaceous","volume":"361","author":"Johnson","year":"2002","journal-title":"Geological Society of America Special Paper"},{"key":"2026061716492711000_CIT0022","doi-asserted-by":"crossref","first-page":"2379","DOI":"10.1126\/science.1072102","article-title":"A tropical rainforest in Colorado 1.4 million years after the Cretaceous-Tertiary boundary","volume":"296","author":"Johnson","year":"2002","journal-title":"Science"},{"key":"2026061716492711000_CIT0023","volume-title":"Plant systematics: a phylogenetic approach","author":"Judd","year":"2018","edition":"4th edn"},{"key":"2026061716492711000_CIT0024","first-page":"133","article-title":"Revision of the flora of the Green River Formation with descriptions of new species","volume":"131F","author":"Knowlton","year":"1923","journal-title":"U.S. Geological Survey Professional Paper"},{"key":"2026061716492711000_CIT0025","first-page":"1","article-title":"The Eocene Green River flora of northwestern Colorado and northeastern Utah","volume":"83","author":"MacGinitie","year":"1969","journal-title":"University of California Publications in Geological Sciences"},{"key":"2026061716492711000_CIT0051","first-page":"1","article-title":"An early middle Eocene flora from the Yellowstone-Absaroka volcanic province, northwestern Wind River Basin, Wyoming","volume":"108","author":"MacGinitie","year":"1974","journal-title":"University of California Publications in Geological Sciences"},{"key":"2026061716492711000_CIT0055","doi-asserted-by":"crossref","first-page":"437","DOI":"10.1111\/nph.13264","article-title":"A metacalibrated time-tree documents the early rise of flowering plant phylogenetic diversity","volume":"207","author":"Magall\u00f3n","year":"2015","journal-title":"New Phytologist"},{"key":"2026061716492711000_CIT0026","doi-asserted-by":"crossref","first-page":"256","DOI":"10.1002\/j.1537-2197.1989.tb11309.x","article-title":"Attached reproductive and vegetative remains of the extinct American-European genus Cedrelospermum (Ulmaceae) from the early Tertiary of Utah and Colorado, USA","volume":"76","author":"Manchester","year":"1989","journal-title":"American Journal of Botany"},{"key":"2026061716492711000_CIT0027","doi-asserted-by":"crossref","first-page":"357","DOI":"10.1086\/719948","article-title":"Extinct anacardiaceous samaras and sumac-like leaves from the Eocene of western North America","volume":"183","author":"Manchester","year":"2022","journal-title":"International Journal of Plant Sciences"},{"key":"2026061716492711000_CIT0028","doi-asserted-by":"crossref","first-page":"897","DOI":"10.1086\/503918","article-title":"Foliage and fruits of early poplars (Salicaceae: Populus) from the Eocene of Utah, Colorado, and Wyoming","volume":"167","author":"Manchester","year":"2006","journal-title":"International Journal of Plant Sciences"},{"key":"2026061716492711000_CIT0046","doi-asserted-by":"crossref","first-page":"281","DOI":"10.2478\/if-2019-0016","article-title":"Bonanzacarpum sprungerorum sp. nov.\u2013a bizarre fruit from the Eocene Green River Formation in Utah, USA","volume":"75","author":"Manchester","year":"2019","journal-title":"Fossil Imprint"},{"key":"2026061716492711000_CIT0030","doi-asserted-by":"crossref","first-page":"53","DOI":"10.18268\/BSGM2014v66n1a5","article-title":"Assembling extinct plants from their isolated parts","volume":"66","author":"Manchester","year":"2014","journal-title":"Bolet\u00edn de la Sociedad Geol\u00f3gica Mexicana"},{"key":"2026061716492711000_CIT0029","doi-asserted-by":"crossref","first-page":"149","DOI":"10.1111\/jse.13011","article-title":"First recognition of the extinct eudicot genus Palibinia in North America: leaves and fruits of Palibinia comptonifolia (RW Br.) comb. nov. from the Eocene of Utah and Colorado, USA","volume":"62","author":"Manchester","year":"2024","journal-title":"Journal of Systematics and Evolution"},{"key":"2026061716492711000_CIT0031","doi-asserted-by":"crossref","first-page":"83","DOI":"10.1007\/s12229-010-9040-1","article-title":"Fossil record and age of the Asteridae","volume":"76","author":"Mart\u00ednez-Mill\u00e1n","year":"2010","journal-title":"The Botanical Review"},{"key":"2026061716492711000_CIT0032","doi-asserted-by":"crossref","first-page":"49","DOI":"10.1186\/s12862-015-0320-6","article-title":"Five major shifts of diversification through the long evolutionary history of Magnoliidae (angiosperms)","volume":"15","author":"Massoni","year":"2015","journal-title":"BMC Evolutionary Biology"},{"key":"2026061716492711000_CIT0052","article-title":"International Code of Nomenclature for algae, fungi, and plants (Melbourne Code)","author":"McNeill","year":"2012","journal-title":"O. bulbotricha"},{"key":"2026061716492711000_CIT0033","doi-asserted-by":"crossref","DOI":"10.1086\/732310","article-title":"Compound-leaved Platanaceae in the Eocene of western North America","volume":"186","author":"Nares","year":"2025","journal-title":"International Journal of Plant Sciences"},{"key":"2026061716492711000_CIT0049","doi-asserted-by":"crossref","first-page":"574","DOI":"10.1666\/07001.1","article-title":"Fossil leaf economics quantified: calibration, Eocene case study, and implications","volume":"33","author":"Royer","year":"2007","journal-title":"Paleobiology"},{"key":"2026061716492711000_CIT0053","doi-asserted-by":"crossref","first-page":"438","DOI":"10.3732\/ajb.0900290","article-title":"Leaf economic traits from fossils support a weedy habit for early angiosperms","volume":"97","author":"Royer","year":"2010","journal-title":"American Journal of Botany"},{"key":"2026061716492711000_CIT0047","doi-asserted-by":"crossref","first-page":"54","DOI":"10.1130\/B26073.1","article-title":"Synoptic reconstruction of a major ancient lake system: Eocene Green River Formation, western United States","author":"Smith","year":"2008","journal-title":"Geological Society of America Bulletin 120"},{"key":"2026061716492711000_CIT0035","doi-asserted-by":"crossref","first-page":"3","DOI":"10.1196\/annals.1438.005","article-title":"Origin and early evolution of angiosperms","volume":"1133","author":"Soltis","year":"2008","journal-title":"Annals of the New York Academy of Sciences"},{"key":"2026061716492711000_CIT0036","first-page":"407","article-title":"Description of the major clades","volume-title":"Phylogeny and evolution of the angiosperms","author":"Soltis","year":"2018"},{"key":"2026061716492711000_CIT0037","doi-asserted-by":"crossref","first-page":"1794","DOI":"10.3732\/ajb.1500298","article-title":"Resolving basal lamiid phylogeny and the circumscription of Icacinaceae with a plastome-scale data set","volume":"102","author":"Stull","year":"2015","journal-title":"American Journal of Botany"},{"key":"2026061716492711000_CIT0050","author":"Stevens","year":"2001"},{"key":"2026061716492711000_CIT0034","first-page":"102","article-title":"A simple device for coating fossils with ammonium chloride","volume":"22","author":"Teichert","year":"1948","journal-title":"Journal of Paleontology"},{"key":"2026061716492711000_CIT0038","doi-asserted-by":"crossref","first-page":"3853","DOI":"10.1073\/pnas.0813376106","article-title":"Rosid radiation and the rapid rise of angiosperm-dominated forests","volume":"106","author":"Wang","year":"2009","journal-title":"Proceedings of the National Academy of Sciences of the United States of America"},{"key":"2026061716492711000_CIT0039","author":"Watson","year":"1992"},{"key":"2026061716492711000_CIT0040","first-page":"1","article-title":"Middle Eocene dicotyledonous plants from Republic, northeastern Washington","volume":"1597","author":"Wolfe","year":"1987","journal-title":"U.S. Geological Survey Bulletin"},{"key":"2026061716492711000_CIT0054","doi-asserted-by":"crossref","first-page":"821","DOI":"10.1038\/nature02403","article-title":"The worldwide leaf economics spectrum","volume":"428","author":"Wright","year":"2004","journal-title":"Nature"},{"key":"2026061716492711000_CIT0041","doi-asserted-by":"crossref","first-page":"416","DOI":"10.54991\/jop.1996.1263","article-title":"Comparison of Miocene fossil floras in lacustrine deposits: implications for palaeoclimatic interpretations at the middle latitudes of the Pacific rim","volume":"45","author":"Yang","year":"1996","journal-title":"Journal of Palaeosciences"}],"container-title":["Annals of Botany"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/academic.oup.com\/aob\/advance-article-pdf\/doi\/10.1093\/aob\/mcae196\/60568551\/mcae196.pdf","content-type":"application\/pdf","content-version":"am","intended-application":"syndication"},{"URL":"https:\/\/academic.oup.com\/aob\/article-pdf\/137\/6\/1971\/60568551\/mcae196.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"syndication"},{"URL":"https:\/\/academic.oup.com\/aob\/article-pdf\/137\/6\/1971\/60568551\/mcae196.pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,6,17]],"date-time":"2026-06-17T20:49:38Z","timestamp":1781729378000},"score":1,"resource":{"primary":{"URL":"https:\/\/academic.oup.com\/aob\/article\/137\/6\/1971\/7888709"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2024,11,9]]},"references-count":53,"journal-issue":{"issue":"6","published-online":{"date-parts":[[2024,11,9]]},"published-print":{"date-parts":[[2026,6,17]]}},"URL":"https:\/\/doi.org\/10.1093\/aob\/mcae196","relation":{},"ISSN":["1095-8290"],"issn-type":[{"value":"1095-8290","type":"electronic"}],"subject":[],"published-other":{"date-parts":[[2026,6]]},"published":{"date-parts":[[2024,11,9]]}}}