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Flora Malesiana Precursor for the Treatment of Moraceae 6: Ficus Subgenus Sycomorus

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Flora Malesiana Precursor for the Treatment of Moraceae 6: Ficus Subgenus Sycomorus BLUMEA 49: 155 –200 Published on 3 May 2004 doi: 10.3767/000651904X486278 FLORA MALESIANA PRECURSOR FOR THE TREATMENT OF MORACEAE 6: FICUS SUBGENUS SYCOMORUS C.C. BERG The Norwegian Arboretum/Botanical Institute, University of Bergen, N-5259 Hjellestad, Norway; Nationaal Herbarium Nederland, Universiteit Leiden branch, P.O. Box 9514, 2300 RA Leiden, The Netherlands SUMMARY The sections of Ficus subg. Sycomorus are described and their Malesian species listed and keyed out. Six new species are described in the subgenus: Ficus albomaculata, F. biakensis, F. boanensis, F. limosa, F. manuselensis, F. morobensis, F. remifolia, F. rubrosyce, F. scopulifera; and one new subspecies: F. botryocarpa Miq. subsp. hirtella C.C. Berg. The new combinations F. botryocarpa Miq. subsp. subalbidoramea (Elmer) C.C. Berg, F. porrecta (Corner) C.C. Berg, F. trichocerasa Diels subsp. pleioclada (Diels) C.C. Berg are also made. The following new sections and subsection are proposed: Subg. Sycomorus sect. Bosscheria (Teijsm. & de Vriese) C.C. Berg, sect. Dammaropsis (Warb.) C.C. Berg, sect. Papuasyce (Corner) C.C. Berg, and subsect. Neomorphe (King) C.C. Berg. Key words: Moraceae, Ficus subg. Sycomorus, Malesia. INTRODUCTION Ficus subg. Sycomorus (Gasp.) Miq. is described and discussed in Flora Malesia precur- sor 1 (Berg, 2003). This contribution deals with the subdivision of this subgenus, the species currently recognised for the region, the new species and subspecies discovered, and provides a key to the Malesian species. The formal subdivision is limited to sections in which a number of informal groups of presumably related species are distinguished; the ranks of series and subseries are not applied. Most of the varieties recognised by Corner (1960a–d, 1961, 1962, 1965) are not maintained, some are recognized as species, some others transferred to other species, as indicated in the lists of Malesian species. SUBDIVISIONS AND SPECIES Proposing a subdivision of subg. Sycomorus is less easy than for the other subgenera. The majority of the species can be accommodated without doubt in the major sections Adenosperma, Sycocarpus, and Sycomorus, as well as in the smaller sections Dam­ maropsis and Papuasyce. An important differentiating floral character used is found in the perianth of the pistillate flower: the tepals are free or nearly so or the tepals are fused, entirely or largely so. In a few species this distinction is unclear. Ficus auriculata comprises individuals of which the tepals are entirely connate and others in which they are free. This has created some confusion with regard to the position of F. oligodon, currently included in F. auriculata (see Corner, 1960b: 39; 1962: 395; © 2004 Nationaal Herbarium Nederland, Leiden University branch 156 BLUMEA — Vol. 49, No. 1, 2004 1978: 383). The other problematical species is F. indigofera Rech. (from the Solomon Islands) showing a considerable variation in the degree of connation of the tepals (see Corner, 1967: 135). Corner (1960a: 418; 1969: 321) placed F. pseudopalma in (his) section Ficus because of the free tepals of the pistillate flowers, but also the related species F. rivularis, in spite of the presence of connate tepals. Evident relatives of F. pseudopalma, currently accommodated in section Dammaropsis, have largely connate tepals. Another problem is linked to several Melanesian species. Ficus cynaroides Corner, F. immanis Corner, and F. lancibracteata Corner, were placed by Corner (1960d: 38; 1967: 122) in subsect. Auriculisperma ser. Cynaroides (sect. Sycocarpus). These three species resemble species of sect. Sycocarpus subsect. Sycocarpus except in the shape of the fruits, and probably also in the absence of internal hairs and hairs on the styles, both usually present in subsect. Sycocarpus. The situation with regard to F. indigofera and F. vitiensis Seem. (from Fiji) is similar. Corner (1960d: 38; 1967: 123) placed them in ser. Vitiensis of subsect. Auriculisperma (sect. Sycocarpus). In particular F. indigofera resembles even in the presence of Termi­ nalia-branching species placed in sect. Adenosperma, but it differs in the presence of entirely connate tepals of pistillate flowers and probably also in the absence of internal hairs. These two sets of species are distinct in having species of Ceratosolen subg. Strepitus as pollinators and not species the subg. Rothropus, found in the majority of the spe- cies of subsect. Sycocarpus, or of subg. Ceratosolen, found in all the species of sect. Adenosperma and some species of subsect. Sycocarpus. Species of subg. Strepitus are also involved in the pollination of species of the sections Dammaropsis and Papuasyce (Wiebes, 1994). In spite of the morphological differences indicated and the different groups of pol- linators involved, F. cynaroides and its relatives and F. indigofera and its relative are currently included in subsect. Sycocarpus and sect. Adenosperma, respectively. The auriculiform to lenticular fruits without the prominent pseudohilum, characteris- tic of subsect. Sycocarpus, and without double keel characteristic of sect. Adenosperma are also found in the two rheophytic species included in subsect. Macrostyla (of sect. Sycocarpus). It is curious that the various small groups of species are so diverse but share the same type of fruit and the same group of pollinators with the possible exception of the Macrostyla species for which the pollinators are not known. The position of the three species placed in sect. Hemicardia and the two placed in sect. Bosscheria is somewhat problematical as is indicated below. SURVEY OF THE SUBDIVISIONS The numbers in parenthesis give the total numbers of species, followed by the numbers of Malesian species. Subg. Sycomorus (c. 135 : 102) Sect. Sycomorus (19 : 6) Subsect. Sycomorus (13 : 1) Subsect. Neomorphe (6 : 5) C. C. Berg: Flora Malesiana precursor for the treatment of Moraceae 6 157 Sect. Hemicardia (3 : 1) Sect. Adenosperma (19 : 16) Sect. Dammaropsis (5 : 3) Sect. Papuasyce (3 : 2) Sect. Bosscheria (2 : 2) Sect. Sycocarpus (86 : 73) Subsect. Sycocarpus (84 : 72) ‘Axillares’ F. calcarata-group F. lepicarpa-group ‘Flagelliflorae’ F. geocarpa-group F. ribes-group F. stolonifera-group F. subterranea-group ‘Cauliflorae’ F. pachyrrhachis-group F. congesta-group Mixed F. cereicarpa-group Subsect. Macrostyla (2 : 1) Ficus subg. Sycomorus (Gasp.) Miq. (1867b) 294 sect. Sycomorus (Gasp.) Miq. (1859) 319 Trees, monoecious or dioecious; internodes not distinctly different in length and leaves not tufted. Leaves spirally arranged and lamina symmetric or distichous and lamina asymmetric, lamina (sub)coriaceous to chartaceous; cystoliths only beneath; waxy glands in the axils of the basal lateral veins or additional ones elsewhere on the lamina beneath (or on the nodes of leafy twigs); petiole rather long. Figs cauliflorous, flagelliflorous (geocarpic) or axillary; basal bracts 3, verticillate; lateral bracts absent. Staminate flowers subtended by 2 bracteoles; stamens (1 or) 2 (or 3). Tepals of pistil- late flowers (2–)3–6, free or connate, often irregular in shape, laciniate and/or narrow, glabrous; styles of long-styled flowers glabrous (or hairy). Fruits lenticular, slightly (simply) keeled, smooth or ± tuberculate, red-brown (or whitish). Distribution — The section comprises 18 species and ranges from West Africa to Australia and the Solomon Islands. Morphology — The section comprises both monoecious and dioecious species. The two groups of species do not differ conspicuously in other characters, neither of vegetative parts nor of flowers and fruits. Subdivision — The dioecious and monoecious species are ranked in different sub- sections. Note — Covellia Gasp. (1844) was based on Ficus ulmifolia Lam. (= C. ulmifolia (Lam.) Gasp.), a species of subg. Sycidium sect. Sycidium in the synonymy of which Covellia is to be included. In 1845, Gasparrini also included F. oppositifolia Willd. in Covellia, currently in the synonymy of F. hispida L.f. which is a species of subg. Sycomorus sect. Sycocarpus. The majority of the Covellia species described by Miquel 158 BLUMEA — Vol. 49, No. 1, 2004 (1848) belong to subg. Sycomorus, in particular to sect. Sycocarpus, to which Corner (1960d) referred Covellia. Ficus subg. Sycomorus sect. Sycomorus subsect. Sycomorus Trees monoecious. Nodal waxy glands absent. Figs without internal hairs. Styles gla- brous. Fruits smooth, brownish. Distribution — Africa to Yemen, Madagascar (and adjacent islands) with 12 species, and F. racemosa from Sri Lanka to Australia. Morphology — The section is rather diverse in the African region, as with regard to the size of the trees, the position and dimensions of the figs (see Berg & Wiebes, 1992, 73–84). Ficus racemosa is morphologically close to the widespread African F. sur Forssk. The Malesian species of the subsection: 1) F. racemosa L. Ficus subg. Sycomorus sect. Sycomorus subsect. Neomorphe (King) C.C. Berg Based on: Ficus sect. Neomorphe King (1887) 2. — Type: Ficus variegata Blume. Trees dioecious. Nodal waxy glands absent or present. Figs with or without internal hairs. Styles mostly without hairs. Fruits ± tuberculate, whitish to brown. Distribution — From Pakistan to Australia and the Solomon Islands; with the excep- tion of F. hainanensis Merr. & Chun, known from China, Indochina, and Thailand, all of them are found in Malesia. Delimitation — The subsection comprises also Tremotis Raf. (1838) 58; Ficus sect. Tremotis (Raf.) Kuntze in Von Post & Kuntze (1904) 236; Ficus subg. Ficus sect. Syco­ carpus subsect. Pomifera
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