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A Revision of Section Urostigma See discussions, stats, and author profiles for this publication at: https://www.researchgate.net/publication/263494526 A Revision of Ficus Subsection Urostigma (Moraceae) Article in Systematic Botany · June 2013 DOI: 10.1600/036364413X670241 CITATIONS READS 6 752 1 author: Bhanumas Chantarasuwan Natural History Museum, Thailand 16 PUBLICATIONS 224 CITATIONS SEE PROFILE Some of the authors of this publication are also working on these related projects: Diversity of Plants and Animals in Thung Yai Naresuan Wildlife Sanctuary, Tak Province View project All content following this page was uploaded by Bhanumas Chantarasuwan on 27 August 2015. The user has requested enhancement of the downloaded file. A Revision of Ficus Subsection Urostigma (Moraceae) Author(s): Bhanumas Chantarasuwan , Cornelis C. Berg , and Peter C. van Welzen Source: Systematic Botany, 38(3):653-686. 2013. Published By: The American Society of Plant Taxonomists URL: http://www.bioone.org/doi/full/10.1600/036364413X670241 BioOne (www.bioone.org) is a nonprofit, online aggregation of core research in the biological, ecological, and environmental sciences. BioOne provides a sustainable online platform for over 170 journals and books published by nonprofit societies, associations, museums, institutions, and presses. Your use of this PDF, the BioOne Web site, and all posted and associated content indicates your acceptance of BioOne’s Terms of Use, available at www.bioone.org/page/terms_of_use. Usage of BioOne content is strictly limited to personal, educational, and non-commercial use. Commercial inquiries or rights and permissions requests should be directed to the individual publisher as copyright holder. BioOne sees sustainable scholarly publishing as an inherently collaborative enterprise connecting authors, nonprofit publishers, academic institutions, research libraries, and research funders in the common goal of maximizing access to critical research. Systematic Botany (2013), 38(3): pp. 653–686 © Copyright 2013 by the American Society of Plant Taxonomists DOI 10.1600/036364413X670241 A Revision of Ficus Subsection Urostigma (Moraceae) Bhanumas Chantarasuwan,1,2,3 Cornelis C. Berg,1 and Peter C. van Welzen1 1Naturalis Biodiversity Center (section NHN), Leiden University, Post Office Box 9514, 2300 RA Leiden, The Netherlands. 2Thailand Natural History Museum, National Science Museum, Klong 5, Klong Luang, Pathum Thani, 12120, Thailand. 3Author for correspondence ([email protected]) Communicating Editor: Fernando Zuloaga Abstract—The present taxonomic revision of Ficus subsection Urostigma recognizes 27 species, of which three are new: F. chiangraiensis, F. middletonii, F. pseudoconcinna. Two new varieties are distinguished within F. virens, var. dispersa and var. matthewii. Ficus lecardii and L. salicifolia, formerly subspecies of F. cordata, are again reinstated to the species level. Typical characters for the subsection are monoecy, monostaminate flowers, red(–brown) colored ovaries and cystoliths on only the abaxial leaf surface. Ficus amplissima and F. rumphii (section Leucogyne) were formerly part of subsection Urostigma, and they have been added here to the key and descriptions because of their morphological resemblance with the species in F. subsection Urostigma. Molecular based phylogenetic analyses showed that at least F. rumphii is unrelated to subsection Urostigma. The two species only differ from subsect. Urostigma in their whitish ovaries and cystoliths at both sides of the leaf blade and they are pollinated by a different group of wasps, species of Eupristina subg. Parapristina. Keywords—Monoecy, morphology, section Leucogyne. Presently, Ficus L. subsect. Urostigma (Gasp.) Berg contains Berg (1989) recognized two main groups in Ficus accord- 27 species, of which seven species are from continental ing to morphological and functional traits, in particular in Africa, Madagascar, and the Arabian Peninsula; and 20 spe- connection to the unique pollination system by fig wasps. cies from Asia, Australia, and the Pacific. Typical are the The first group comprises the subgenera Pharmacosycea and tree habit, intermittent growth, often deciduous, leaves Urostigma, and the other contains the subgenera Ficus, spirally arranged, often articulate or subarticulate, cystoliths Sycidium, and Sycomorus. Fourteen years later, Berg (2003) only abaxially, figs axillary, more commonly just below divided Ficus into six subgenera (Pharmacosycea, Urostigma, the leaves, and/or ramiflorous on up to ca. 1 cm long spurs, Ficus, Synoecia, Sycidium, and Sycomorus) based on major staminate flowers near the ostiole or scattered among the differentiating characters like monoecy–dioecy, adventitious pistillate ones, tepals red(dish), ovary red–brown (or white). roots, stipules, position of figs, bracts, stigmas, and waxy This taxon started in a less elaborate circumscription as glands. This new classification did not completely correlate Ficus subgenus Urostigma, which was first described by any longer with the pollination system; exceptions occurred. Miquel in 1867. Miquel abandoned the idea of breaking One exception was Berg’s (2004) newly established sub- up the genus Ficus into genera and he divided Ficus into section Urostigma.Berg(2004)unitedCorner’ssections six subgenera. Urostigma is one of the subgenera, then Leucogyne, Urostigma,andConosycea,intosect.Urostigma, comprising ca. 270 species. Urostigma contains monoecious which he subdivided into subsect. Urostigma (containing species with unistaminate or bistaminate flowers. How- Corner’s sect. Leucogyne and Urostigma) and subsect. Conosycea ever, Miquel subdivided subgen. Urostigma according to (containing Corner’s sect. Conosycea). Typical for subsect. distribution, recognizing six series for Asia and Australia, Urostigma is the presence of intermittent growth. Berg did three series for Africa, and five series for America. The not recognize the series used by Corner. Berg also included species presently included in subsect. Urostigma were mainly African species in the otherwise Asian/Pacific subsection. placed in the series Infectoriae Miq. and Religiosae Miq. of Asia The two species in former section Leucogyne are pollinated and Australia. The African representatives of the subsection by different wasps than the rest of subsect. Urostigma (see were classified in the series Grandiores Miq., Oblongifoliae paragraph on pollination below). Miq., and Ellipticifoliae Miq. Berg and Corner (2005) remarked that the differences In 1887 King divided Ficus into seven sections of which between Corner’s sect. Leucogyne and Urostigma are not good sect. Urostigma was equivalent to Miquel’s subgenus. King enough to justify recognition on a sectional or subsectional divided his section Urostigma into series and subseries taxonomic level, and thus subsection Urostigma containing based on leaf characters. The species of the present treat- both taxa was maintained. ment occur in different subseries of King. Recently, results of molecular phylogenetic studies by In 1960, Corner reused the rank “subgenus.” He recog- Rønsted et al. (2005, 2008) show that F. rumphii is not part nized three subgenera, one of them subg. Urostigma with of subsect. Urostigma, but that it is embedded in the not seven sections, among which sect. Urostigma (with 15 spe- closely related subsection Conosycea. Unfortunately, Rønsted cies in Asia and six species in Africa) and sect. Leucogyne et al. (2005, 2008) did not sample F. amplissima, the other with two species in Asia (F. amplissima J. E. Sm. and species in former sect. Leucogyne, which is thus phylo- F. rumphii Blume). These two groups contain the species genetically still incompletely known. treated here. The main difference between the two sections Transferring both species to subsection Conosycea com- are the color of the ovary, the position of the staminate plicates the morphological distinctiveness of the subsections flowers and the position of the cystoliths: whitish ovaries, as the two species share major characters with subsect. staminate flowers dispersed and cystoliths at both sides Urostigma: (1) Both species show an intermittent growth like of the leaf blade in Leucogyne, and red(–brown) ovaries, in subsect. Urostigma. (2) Staminate flowers of F. amplissima staminate flowers around the ostiole (not 100%) and cysto- and F. rumphii are completely dispersed throughout the liths only abaxially in Urostigma. fig, which is also present in F. orthoneura H. Le´v. & Vaniot, 653 654 SYSTEMATIC BOTANY [Volume 38 F. hookeriana Corner, and F. prolixa G. Forst. of subsect. glabrous on both sides, but the young leaves of F. cupulata Urostigma,whileF. arnottiana (Miq.) Miq. and F. virens Aiton Haines are sometimes puberulous. The margin is always var. dispersa Chantaras. show a transition with abundant entire. The venation is basically pinnate and brochidodromous. staminate flowers around the ostiole and a few dispersed The basal lateral veins are distinct by the narrower angle ones. (3) Leaf articulation is absent in subsect. Conosycea, of departure from the midrib. Some species, e.g. F. rumphii, section Leucogyne and in the African and Malagasy species F. ingens (Miq.) Miq., and F. cupulata, have branched basal of subsect. Urostigma, but mostly present in the Asian spe- veins. The tertiary venation varies from clearly scalariform cies of subsect. Urostigma (F. orthoneura and F. hookeriana to reticulate and/or partly parallel to the lateral veins. The excepted). (4) Color of the ovary is normally
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