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A Revision of Xylopia L. (Annonaceae): the Species of Madagascar and the Mascarene Islands

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A Revision of Xylopia L. (Annonaceae): the Species of Madagascar and the Mascarene Islands adansonia 2020 ● 42 ● 1 DIRECTEUR DE LA PUBLICATION : Bruno David Président du Muséum national d’Histoire naturelle RÉDACTEUR EN CHEF / EDITOR-IN-CHIEF : Thierry Deroin RÉDACTEURS / EDITORS : Porter P. Lowry II ; Zachary S. Rogers ASSISTANTS DE RÉDACTION / ASSISTANT EDITORS : Emmanuel Côtez ([email protected]) ; Anne Mabille MISE EN PAGE / PAGE LAYOUT : Emmanuel Côtez COMITÉ SCIENTIFIQUE / SCIENTIFIC BOARD : P. Baas (Nationaal Herbarium Nederland, Wageningen) F. Blasco (CNRS, Toulouse) M. W. Callmander (Conservatoire et Jardin botaniques de la Ville de Genève) J. A. Doyle (University of California, Davis) P. K. Endress (Institute of Systematic Botany, Zürich) P. Feldmann (Cirad, Montpellier) L. Gautier (Conservatoire et Jardins botaniques de la Ville de Genève) F. Ghahremaninejad (Kharazmi University, Téhéran) K. Iwatsuki (Museum of Nature and Human Activities, Hyogo) K. Kubitzki (Institut für Allgemeine Botanik, Hamburg) J.-Y. Lesouef (Conservatoire botanique de Brest) P. Morat (Muséum national d’Histoire naturelle, Paris) J. Munzinger (Institut de Recherche pour le Développement, Montpellier) S. E. Rakotoarisoa (Millenium Seed Bank, Royal Botanic Gardens Kew, Madagascar Conservation Centre, Antananarivo) É. A. Rakotobe (Centre d’Applications des Recherches pharmaceutiques, Antananarivo) P. H. Raven (Missouri Botanical Garden, St. Louis) G. Tohmé (Conseil national de la Recherche scientifique Liban, Beyrouth) J. G. West (Australian National Herbarium, Canberra) J. R. Wood (Oxford) COUVERTURE / COVER : Made from the Figures of the article. Adansonia est indexé dans / Adansonia is indexed in: – Science Citation Index Expanded (SciSearch®) – ISI Alerting Services® – Current Contents® / Agriculture, Biology, and Environmental Sciences® – Scopus® Adansonia est distribué en version électronique par / Adansonia is distributed electronically by: – BioOne® (http://www.bioone.org) Adansonia est une revue en flux continu publiée par les Publications scientifiques du Muséum, Paris Adansonia is a fast track journal published by the Museum Science Press, Paris Les Publications scientifiques du Muséum publient aussi / The Museum Science Press also publish: Geodiversitas, Zoosystema, Anthropozoologica, European Journal of Taxonomy, Naturae, Cryptogamie sous-sections Algologie, Bryologie, Mycologie. Diffusion – Publications scientifiques Muséum national d’Histoire naturelle CP 41 – 57 rue Cuvier F-75231 Paris cedex 05 (France) Tél. : 33 (0)1 40 79 48 05 / Fax : 33 (0)1 40 79 38 40 [email protected] / http://sciencepress.mnhn.fr © Publications scientifiques du Muséum national d’Histoire naturelle, Paris, 2019 ISSN (imprimé / print) : 1280-8571/ ISSN (électronique / electronic) : 1639-4798 A revision of Xylopia L. (Annonaceae): the species of Madagascar and the Mascarene islands David M. JOHNSON Department of Botany and Microbiology Ohio Wesleyan University Delaware, Ohio 43015 (United States) [email protected] Nancy A. MURRAY Department of Botany and Microbiology Ohio Wesleyan University Delaware, Ohio 43015 (United States) [email protected] Submitted on 2 March 2019 | accepted on 2 July 2019 | Published on 5 February 2020 Johnson D. M. & Murray N. A. 2020. — A revision of Xylopia L. (Annonaceae): the species of Madagascar and the Mas- carene islands. Adansonia, sér. 3, 42 (1): 1-88. https://doi.org/10.5252/adansonia2020v42a1. http://adansonia.com/42/1 ABSTRACT We recognize 30 species of the pantropical genus Xylopia L. from Madagascar and an additional three species from the Mascarene Islands. Of the six sections of the genus, three are represented in the region: sect. Xylopia, sect. Verdcourtia, and sect. Stenoxylopia. All species are endemic, and many KEY WORDS are microendemics of conservation concern. The greatest species richness occurs in humid forests Xylopia, below 1000 m. Taxonomic review supports the hypothesis that the genus dispersed to Madagascar pantropical Annonaceae, Madagascar, from continental Africa five times. Ten new species are described:Xylopia anomala D.M.Johnson & Mascarene, N.A.Murray, sp. nov., X. australis D.M.Johnson & N.A.Murray, sp. nov., X. carinata D.M.Johnson & islands, long-distance dispersal, N.A.Murray, sp. nov., X. galokothamna D.M.Johnson & N.A.Murray, sp. nov., X. lokobensis bird/lemur dispersal, D.M.Johnson & N.A.Murray, sp. nov., X. longirostra D.M.Johnson & N.A.Murray, sp. nov., X. maro- microendemism, jejyana D.M.Johnson & N.A.Murray, sp. nov., X. ravelonarivoi D.M.Johnson & N.A.Murray, sp. nov., conservation, lectotypification, X. retusa D.M.Johnson & N.A.Murray, sp. nov., and X. sclerophylla D.M.Johnson & N.A.Murray, new species. sp. nov. An identification key, technical descriptions, illustrations, and distribution maps are provided. RÉSUMÉ Révision du genre Xylopia L. (Annonaceae) : les espèces de Madagascar et des îles Mascareignes. Nous reconnaissons 30 espèces du genre pantropical Xylopia L. qui viennent de Madagascar et trois espèces additionnelles des Mascareignes. Des six sections du genre, trois sont représentées dans la région : sect. Xylopia, sect. Verdcourtia, et sect. Stenoxylopia. Toutes les espèces sont endémiques, et MOTS CLÉS Xylopia, beaucoup sont des espèces micro-endémiques dont l’état de conservation est préoccupant. La plus Annonaceae grande richesse spécifique se trouve dans les forêts humides en dessous de 1000 m. La révision tax- pantropicales, onomique soutient l’hypothèse que le genre s’est dispersé à Madagascar depuis l’Afrique continentale cinq Madagascar, Mascareignes, fois. Dix nouvelles espèces sont décrites : Xylopia anomala D.M.Johnson & N.A.Murray, sp. nov., X. aus- dispersion à longue- tralis D.M.Johnson & N.A.Murray, sp. nov., X. carinata D.M.Johnson & N.A.Murray, sp. nov., distance, dispersion par oiseaux/ X. galokothamna D.M.Johnson & N.A.Murray, sp. nov., X. lokobensis D.M.Johnson & N.A.Murray, lémuriens, sp. nov., X. longirostra D.M.Johnson & N.A.Murray, sp. nov., X. marojejyana D.M.Johnson & micro-endémisme, N.A.Murray, sp. nov., X. ravelonarivoi D.M.Johnson & N.A.Murray, sp. nov., X. retusa D.M.Johnson & conservation, lectotypification, N.A.Murray, sp. nov. et X. sclerophylla D.M.Johnson & N.A.Murray, sp. nov. Une clé de détermination, espèces nouvelles. des descriptions techniques, des illustrations, et des cartes de distribution sont fournies. ADANSONIA, sér. 3 • 2020 • 42 (1) © Publications scientifiques du Muséum national d’Histoire naturelle, Paris. www.adansonia.com 1 Johnson D. M. & Murray N. A. INTRODUCTION The generic concept ofXylopia was clarified during the first half of the 19th century, and Baillon (1864) worked from this Within the early-divergent angiosperm family Annonaceae improved understanding to review the known African species the genus Xylopia L. is unique in its pantropical distribution of Xylopia, including plants from Madagascar and the Mas- and, with nearly 200 species of trees, shrubs, and lianas, is the carene islands (summarized in Johnson & Murray 2018). He second largest genus in the family. The plants bear distinctive pointed out that Annona grandiflora and A. amplexicaulis had fruits composed of clusters of monocarps that dehisce when ripe, numerous ovules in each carpel, and that their carpels were displaying the seeds against an endocarp of contrasting color not fused at the base but were surrounded by a cuplike sac. (Johnson & Murray 2018). Twenty-five endemic species had He stressed that these characteristics were discordant with previously been described from the floras of Madagascar and Annona but much like those of Xylopia, and accordingly the Mascarene Islands of La Réunion and Mauritius (Cavaco & moved them to the latter genus, although he segregated them Keraudren 1958; Keraudren-Aymonin 1980; Callmander et al. in a distinct section he named Pseudanona. He renamed An- 2009), many from only one or two collections. nona grandiflora as Xylopia lamarckii Baill. because the ep- Xylopia arose in continental Africa and diversified into six ithet grandiflora was preoccupied in Xylopia, described two evolutionary lines that we recognize as sections (Stull et al. new species from Madagascar, X. buxifolia Baill. and X. lastel- 2017, Johnson & Murray 2018). A sample of 19 accessions liana Baill., and described one additional species, X. richardii from Madagascar and the island of Mauritius analyzed in the Boivin ex Baill., from the island of La Réunion. By Baillon’s Stull et al. (2017) study showed the Madascar and Mascarene accounting, there were now four Xylopia species known samples to represent three of the five evolutionary lines present from Madagascar (X. amplexicaulis (Lam.) Baill., X. buxi- in Africa, and the evidence that there have been at least five folia, X. lamarckii, and X. lastelliana), one from Mauritius separate colonizations of the islands from Africa. Eight of the (X. lamarckii), and two from La Réunion (X. lamarckii and 19 Madagascar accessions could not be identified as a known X. richardii). Baker (1877) did not accept Baillon’s transfer species at that time. In order to clarify the taxonomy of the of A. grandiflora and A. amplexicaulis to Xylopia, but valid- Madagascar and Mascarene species of Xylopia, we conducted ly published the name An[n]ona pyriformis Bojer ex Baker. the following revisionary study. In 1882 Baillon described Xylopia humblotiana Baill. from eastern Madagascar. At the beginning of the 20th century Safford (1913) chose to stress the distinctive large fleshy TAXONOMIC HISTORY petals of X. lamarckii and X. amplexicaulis in raising Xylopia section Pseudanona to the level of a distinct
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