J Syst Evol

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Large-scale nuclear and plastid phylogenomic analyses inform an updated Angiosperm Phylogeny Group classification: APG V

James W. Byng1, Mark W. Chase2*, Maarten J. M. Christenhusz1,3, Michael F. Fay2, De-Zhu Li4,5, Hong Ma6, David J. Mabberley7,8,9, Douglas E. Soltis10,11,12,13, Pamela S. Soltis11,12,13, Peter F. Stevens14, William J. Baker2,15, Steven Dodsworth3, Félix Forest2, Olivier Maurin2,16,17, Lisa Pokorny18, Stephen A. Smith19, Alexandre R. Zuntini2   

  1. 1 Hortus Botanicus, Faculty of Architecture and the Built Environment, University of Technology, Delft, The Netherlands;
    2 Royal Botanic Gardens, Kew, Richmond, TW9 3AE, UK;
    3 Institute of Structural and Molecular Biology, School of Natural Sciences, Birkbeck, University of London, London WC1E 7HX, UK;
    4 Center for Interdisciplinary Biodiversity Research and College of Forestry, Shandong Agricultural University, Tai'an 271018, China;
    5 Germplasm Bank of Wild Species and Yunnan Key Laboratory of Crop Wild Relative Omics, Kunming Institute of Botany, Chinese Academy of Sciences, Kunming 650201, China;
    6 Department of Biology, Eberly College of Science and Huck Institutes of the Life Sciences, Pennsylvania State University, University Park, PA, USA;
    7 Wadham College, University of Oxford, Oxford, UK;
    8 School of Natural Resources, Macquarie University, Sydney, New South Wales, Australia;
    9 Australian Institute of Botanical Science (National Herbarium of New South Wales), Sydney, New South Wales, Australia;
    10 Department of Biology, University of Florida, Gainesville, FL 32611, USA;
    11 Florida Museum of Natural History, University of Florida, Gainesville, FL 32611, USA;
    12 Genetics Institute, University of Florida, Gainesville, FL 32611, USA;
    13 Biodiversity Institute, University of Florida, Gainesville, FL 32611, USA;
    14 Missouri Botanical Garden, PO Box 299, St Louis, MO 63166-0299, USA;
    15 Department of Biology, Aarhus University, Aarhus, Denmark;
    16 Meise Botanic Garden, 1860, Meise, Belgium;
    17 Fédération Wallonie-Bruxelles, Service Général de l'Enseignement supérieur et de la Recherche scientifique, 1080, Bruxelles, Belgium;
    18 Real Jardín Botánico (RJB-CSIC), 28014 Madrid, Spain;
    19 Department of Ecology & Evolutionary Biology, University of Michigan, 830 North University Avenue, Ann Arbor, MI, 48109-1048 USA
    *Author for correspondence. E-mail: m.chase@kew.org
  • Received:2025-12-11

Abstract: We present here a revision of the APG classification that considers the extensive recent analyses of hundreds of nuclear and plastid genes for many angiosperm species. Although previous versions of the APG classification were largely based on uniparentally inherited markers (plastid DNA, largely maternally inherited), there has emerged since APG IV extensive evidence of widespread hybridization and inheritance of ancestral polymorphisms. Despite this evidence of gene-tree discordance, most of the APG IV classification is supported by both plastid and nuclear analyses, and only a few revisions are required in this update to make the classification parallel phylogenetic results in the literature. At the ordinal level, few changes in circumscription are required relative to the last APG version, although the fabids now comprise only the four nitrogen-fixing orders (Cucurbitales, Fabales, Fagales, and Rosales) and the malvids now include the former COM clade (Celastrales, Malpighiales, and Oxalidales), which in nuclear DNA analyses is no longer monophyletic. Oncothecales (with only Oncothecaceae) and Cardiopteridales (with Cardiopteridaceae and Stemonuraceae) are newly recognized, and Icacinales are restricted to Icacinaceae. Restriction of Aquifoliales and Bruniales to just Aquifoliaceae/Helwingiaceae and Bruniaceae, respectively, is also proposed. Huaceae and Columelliaceae are unplaced to order among the malvids and campanulids, respectively. At the family level, expanded circumscriptions include Tecophilaeaceae (including Ixioliriaceae, Asparagales), Frankeniaceae (including Tamaricaceae, Caryophyllales), Phytolaccaceae (including Agdestidaceae and Sarcobataceae, Caryophyllales), Helwingiaceae (including Phyllonomaceae, Aquifoliales), Gesneriaceae (including Calceolariaceae and Peltantheraceae, Lamiales), Pentaphylacaceae (including Sladeniaceae, Ericales), and Orobanchaceae (including Mazaceae, Paulowniaceae, Phrymaceae, and Wightiaceae, Lamiales). In Santalales, the limits of Santalaceae and Olacaceae are revised, and Erythropalaceae and Strombosiaceae are recognized as distinct from Olacaceae; Balanophoraceae are maintained as distinct from Santalaceae, but this is a tentative placement. Generic relationships in Dioscoreales require more analyses before a revised family classification can be proposed, so we maintain the APG IV families of this order.