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      Insights on the systematics and morphology of Humiriaceae ( Malpighiales ): androecial and extrafloral nectary variation, two new combinations, and a new Sacoglottis from Guyana

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          Abstract

          Abstract

          Humiriaceae have had little recent comparative morphological study except for their distinctive fruits. We surveyed the diversity of stamen structures in the family with consideration of dehiscence patterns and the evolutionary transitions between tetra- and disporangiate anthers. Novel interpretations of floral morphology support new combinations ( Duckesia liesneri K.Wurdack & C.E.Zartman, comb. nov. and Vantanea spiritu-sancti K.Wurdack & C.E.Zartman, comb. nov.) for two species formerly in Humiriastrum . We investigated all eleven species of Sacoglottis for diagnostic features that may contribute to better species delimitations, and describe Sacoglottis perryi K.Wurdack & C.E.Zartman, sp. nov. as an endemic of the Pakaraima Mountains in western Guyana. Finally, our survey across Humiriaceae for extrafloral nectaries (EFNs) revealed their presence on leaves of all extant species as adaxial basilaminar and/or abaxial embedded glands, in addition to the frequent occurrence of marginal glandular setae. The significance of inter-generic variation in gland position and anther morphology within the family are discussed.

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          Most cited references34

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          The Ecology and Evolution of Ant-Plant Interactions

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            Malpighiales phylogenetics: Gaining ground on one of the most recalcitrant clades in the angiosperm tree of life.

            The eudicot order Malpighiales contains ∼16000 species and is the most poorly resolved large rosid clade. To clarify phylogenetic relationships in the order, we used maximum likelihood, Bayesian, and parsimony analyses of DNA sequence data from 13 gene regions, totaling 15604 bp, and representing all three genomic compartments (i.e., plastid: atpB, matK, ndhF, and rbcL; mitochondrial: ccmB, cob, matR, nad1B-C, nad6, and rps3; and nuclear: 18S rDNA, PHYC, and newly developed low-copy EMB2765). Our sampling of 190 taxa includes representatives from all families of Malpighiales. These data provide greatly increased support for the recent additions of Aneulophus, Bhesa, Centroplacus, Ploiarium, and Rafflesiaceae to Malpighiales; sister relations of Phyllanthaceae + Picrodendraceae, monophyly of Hypericaceae, and polyphyly of Clusiaceae. Oxalidales + Huaceae, followed by Celastrales are successive sisters to Malpighiales. Parasitic Rafflesiaceae, which produce the world's largest flowers, are confirmed as embedded within a paraphyletic Euphorbiaceae. Novel findings show a well-supported placement of Ctenolophonaceae with Erythroxylaceae + Rhizophoraceae, sister-group relationships of Bhesa + Centroplacus, and the exclusion of Medusandra from Malpighiales. New taxonomic circumscriptions include the addition of Bhesa to Centroplacaceae, Medusandra to Peridiscaceae (Saxifragales), Calophyllaceae applied to Clusiaceae subfamily Kielmeyeroideae, Peraceae applied to Euphorbiaceae subfamily Peroideae, and Huaceae included in Oxalidales.
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              Phylogenomics and a posteriori data partitioning resolve the Cretaceous angiosperm radiation Malpighiales.

              The angiosperm order Malpighiales includes ~16,000 species and constitutes up to 40% of the understory tree diversity in tropical rain forests. Despite remarkable progress in angiosperm systematics during the last 20 y, relationships within Malpighiales remain poorly resolved, possibly owing to its rapid rise during the mid-Cretaceous. Using phylogenomic approaches, including analyses of 82 plastid genes from 58 species, we identified 12 additional clades in Malpighiales and substantially increased resolution along the backbone. This greatly improved phylogeny revealed a dynamic history of shifts in net diversification rates across Malpighiales, with bursts of diversification noted in the Barbados cherries (Malpighiaceae), cocas (Erythroxylaceae), and passion flowers (Passifloraceae). We found that commonly used a priori approaches for partitioning concatenated data in maximum likelihood analyses, by gene or by codon position, performed poorly relative to the use of partitions identified a posteriori using a Bayesian mixture model. We also found better branch support in trees inferred from a taxon-rich, data-sparse matrix, which deeply sampled only the phylogenetically critical placeholders, than in trees inferred from a taxon-sparse matrix with little missing data. Although this matrix has more missing data, our a posteriori partitioning strategy reduced the possibility of producing multiple distinct but equally optimal topologies and increased phylogenetic decisiveness, compared with the strategy of partitioning by gene. These approaches are likely to help improve phylogenetic resolution in other poorly resolved major clades of angiosperms and to be more broadly useful in studies across the Tree of Life.
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                Author and article information

                Contributors
                Journal
                PhytoKeys
                PhytoKeys
                3
                urn:lsid:arphahub.com:pub:f7fce910-8e78-573f-9c77-7788555f8aad
                PhytoKeys
                Pensoft Publishers
                1314-2011
                1314-2003
                2019
                20 June 2019
                : 124
                : 87-121
                Affiliations
                [1 ] Department of Botany, MRC-166, National Museum of Natural History, Smithsonian Institution, P.O. Box 37012, Washington, DC 20013-7012, USA National Museum of Natural History, Smithsonian Institution Washington D.C. United States of America
                [2 ] Department of Biodiversity, National Institute for Amazonian Research (INPA), Av. André, Araújo 2936, Aleixo, Manaus, Amazonas 69060-001, Brazil National Institute of Amazonian Research (INPA) and Smithsonian Tropical Research Institute Manaus Brazil
                Author notes
                Corresponding author: Kenneth J. Wurdack ( wurdackk@ 123456si.edu )

                Academic editor: A. Sennikov

                Article
                PMC6598937 PMC6598937 6598937 34679
                10.3897/phytokeys.124.34679
                6598937
                31274979
                a85231a1-b6a4-4d72-b95a-2f16a6404120

                This is an open access article distributed under the terms of the CC0 Public Domain Dedication.

                History
                : 20 March 2019
                : 28 April 2019
                Categories
                Research Article
                Humiriaceae
                Evolutionary Biology
                Taxonomy
                Americas

                Duckesia , Vantanea , Humiriastrum ,glands,floral morphology,extrafloral nectaries,anthers

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