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Ficus Subsection Urostigma (Moraceae) Issue Date: 2014-09-03 Chapter 2 a Revision of Ficus Subsection Urostigma (Moraceae)

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Cover Page The handle http://hdl.handle.net/1887/28481 holds various files of this Leiden University dissertation Author: Chantarasuwan, Bhanumas Title: Taxonomy, systematics, and biogeography of Ficus subsection Urostigma (Moraceae) Issue Date: 2014-09-03 Chapter 2 A revision of Ficus subsection Urostigma (Moraceae) Bhanumas Chantarasuwan, Cornelis C. Berg, and Peter C. van Welzen Published in: Systematic Botany 38(3), 2013: 653-686. 20 21 A REVISION OF FICUS SUBSECTION UROSTIGMA (MORACEAE) Abstract Introduction The present taxonomic revision ofFicus subsection Urostigma recognizes Presently, Ficus L. subsect. Urostigma (Gasp.) Berg contains 27 species, of 27 species, of which three are new: F. chiangraiensis, F. middletonii, which seven species are from continental Africa, Madagascar, and the Arabian F. pseudoconcinna. Two new varieties are distinguished within F. virens, var. Peninsula; and 20 species from Asia, Australia, and the Pacific. Typical are dispersa and var. matthewii. Ficus lecardii and F. salicifolia, formerly subspecies the tree habit, intermittent growth, often deciduous, leaves spirally arranged, of F. cordata, are again reinstated to the species level. Typical characters for often articulate or subarticulate, cystoliths only abaxially, figs axillary, more the subsection are monoecy, monostaminate flowers, red(brown) colored commonly just below the leaves, and/or ramiflorous on up to ca. 1 cm long ovaries and cystoliths on only the abaxial leaf surface. Ficus amplissima and spurs, staminate flowers near the ostiole or scattered among the pistillate ones, F. rumphii (section Leucogyne) were formerly part of subsection Urostigma, tepals red(dish), ovary red brown (or white). and they have been added here to the key and descriptions because of their morphological resemblance with the species in subsection Urostigma. This taxon started in a less elaborate circumscription as Ficus subgenus Molecular–based phylogenetic analyses showed that at least F. rumphii is Urostigma, which was first described by Miquel in 1867. Miquel abandoned unrelated to subsection Urostigma. The two species only differ from subsect. the idea of breaking up the genus Ficus into genera and he divided Ficus Urostigma in their whitish ovaries and cystoliths at both sides of the leaf blade into six subgenera. Urostigma is one of the subgenera, then comprising ca. and they are pollinated by a different group of wasps, species of Eupristina 270 species. Urostigma contains monoecious species with unistaminate subg. Parapristina. or bistaminate flowers. However, Miquel subdivided subgen. Urostigma according to distribution, recognizing six series for Asia and Australia, three Keywords–Monoecy, morphology, section Leucogyne. series for Africa, and five series for America. The species presently included in subsect. Urostigma were mainly placed in the series Infectoriae Miq. and Religiosae Miq. of Asia and Australia. The African representatives of the subsection were classified in the seriesGrandiores Miq., Oblongifoliae Miq., and Ellipticifoliae Miq. In 1887 King divided Ficus into seven sections of which sect. Urostigma was equivalent to Miquel’s subgenus. King divided his section Urostigma into series and subseries based on leaf characters. The species of the present treatment occur in different subseries of King. In 1960, Corner reused the rank “subgenus.” He recognized three subgenera, one of them subg. Urostigma with seven sections, among which sect. Urostigma (with 15 species in Asia and six species in Africa) and sect. Leucogyne with two species in Asia (F. amplissima J. E. Sm. and F. rumphii Blume). These two groups contain the species treated here. The main difference between the two sections are the color of the ovary, the position of the staminate flowers and the position of the cystoliths: whitish ovaries, staminate flowers dispersed and cystoliths at both sides of the leaf blade in Leucogyne, and red(brown) ovaries, staminate flowers around the ostiole (not 100%) and cystoliths only abaxially in Urostigma. Berg (1989) recognized two main groups in Ficus according to morphological and functional traits, in particular in connection to the unique pollination system by fig wasps. The first group comprises the subgeneraPharmacosycea and Urostigma, and the other contains the subgenera Ficus, Sycidium, and Sycomorus. Fourteen years later, Berg (2003) divided Ficus into six subgenera (Pharmacosycea, Urostigma, Ficus, Synoecia, Sycidium, and Sycomorus) based on major differentiating characters like monoecy–dioecy, adventitious roots, stipules, position of figs, bracts, stigmas, and waxy glands. This new classification did not completely correlate any longer with the pollination 22 23 A REVISION OF FICUS SUBSECTION UROSTIGMA (MORACEAE) system; exceptions occurred. One exception was Berg’s (2004) newly because the phylogenetic status of F. amplissima is unknown. In comparison established subsection Urostigma. Berg (2004) united Corner’s sections with Berg and Corner (2005), this paper treats all species together, not only Leucogyne, Urostigma, and Conosycea, into sect. Urostigma, which he the Malesian ones. Descriptions and nomenclature are more complete and subdivided into subsect. Urostigma (containing Corner’s sect. Leucogyne and based on more specimens, and the latest species delimitations are presented. Urostigma) and subsect. Conosycea (containing Corner’s sect. Conosycea). Typical for subsect. Urostigma is the presence of intermittent growth. Berg Results did not recognize the series used by Corner. Berg also included African species in the otherwise Asian/Pacific subsection. The two species in former section Habit—All species are essentially hemi–epiphytic, but without abundant aerial Leucogyne are pollinated by different wasps than the rest of subsect. Urostigma roots. Some species are often terrestrial. Most of the species remain medium– (see paragraph on pollination below). Berg and Corner (2005) remarked that sized trees, rarely taller than 25 m, but some Asian species, like F. caulocarpa the differences between Corner’s sect.Leucogyne and Urostigma are not good (Miq.) Miq., F. superba (Miq.) Miq., and F. virens Aiton often become 30–35 enough to justify recognition on a sectional or subsectional taxonomic level, m tall (Berg and Corner 2005). The African species F. verruculosa Warb. is a and thus subsection Urostigma containing both taxa was maintained. shrub or a treelet, and the Malagasy species F. madagascariensis C. C. Berg is sometimes a shrub (Berg and Wiebes 1992). The trees show intermittent Recently, results of molecular phylogenetic studies by Rønsted et al. (2005, growth, for which morphological indications are different colors of parts of 2008) show that F. rumphii is not part of subsect. Urostigma, but that it is twigs of current or recent growth and of the previous season’s growth. The embedded in the not closely related subsect. Conosycea. Unfortunately, transition is marked by a section with short internodes, which in some species Rønsted et al. (2005, 2008) did not sample F. amplissima, the other species bear persistent coriaceous stipules, forming terminal buds. in former sect. Leucogyne, which is thus phylogenetically still incompletely known. Indumentum—The indumentum consists of unicellular hairs of whitish, yellowish or brown colors which usually occur on leafy twigs, petioles, Transferring both species to subsection Conosycea complicates the stipules, peduncles, and sometimes on the upper ostiolar bracts. Many species morphological distinctiveness of the subsect. as the two species share have translucent hairs on the inner surface of the fig (receptacle) among the major characters with subsect. Urostigma: (1) Both species show an flowers; these are called “internal hairs.” intermittent growth like in subsect. Urostigma. (2) Staminate flowers of F. amplissima and F. rumphii are completely dispersed throughout the fig, Leaves—The leaves are always spirally arranged. The lamina varies from which is also present in F. densifolia Miq., F. hookeriana Corner, F. orthoneura broadest below the middle to broadest above the middle. The lamina is always H. Lév. & Vaniot and F. prolixa G. Forst. of subsect. Urostigma, while symmetrical and ranges from small (up to 10 cm long) to medium–sized F. arnottiana (Miq.) Miq. and F. virens Aiton var. dispersa Chantaras. show (10–20 cm long), but those of F. hookeriana Corner can be up to 25 cm long. a transition with abundant staminate flowers around the ostiole and a few The lamina is subcoriaceous to coriaceous and lacks a hypodermis except dispersed ones. (3) Leaf articulation is absent in subsect. Conosycea, section for F. hookeriana Corner and F. orthoneura H. Lév. & Vaniot, which have a Leucogyne and in the African and Malagasy species of subsect. Urostigma, but well–developed hypodermis on both sides. The lamina is mostly glabrous on mostly present in the Asian species of subsect. Urostigma (F. orthoneura and both sides, but the young leaves of F. cupulata Haines are sometimes puberulous. F. hookeriana excepted). (4) Color of the ovary is normally a good character The margin is always entire. The venation is basically pinnate and to separate the two subsections, as ovaries in subsect. Conosycea are white brochidodromous. The basal lateral veins are distinct by the narrower angle (including the two Leucogyne species) and red(brown) in subsect. Urostigma. of departure from the midrib. Some species, e.g. F. rumphii, F. ingens However, the ovary of F. arnottiana is white
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