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Lecythidaceae (G.T Flora Malesiana, Series I, Volume 21 (2013) 1–118 LECYTHIDACEAE (G.T. Prance, Kew & E.K. Kartawinata, Bogor)1 Lecythidaceae A.Rich. in Bory, Dict. Class. Hist. Nat. 9 (1825) 259 (‘Lécythidées’), nom. cons.; Poit., Mém. Mus. Hist. Nat. Paris 13 (1835) 141; Miers, Trans. Linn. Soc. London, Bot. 30, 2 (1874) 157; Nied. in Engl. & Prantl, Nat. Pflanzenfam. 3, 7 (1892) 30; R.Knuth in Engl., Pflanzenr. IV.219, Heft 105 (1939) 26; Whitmore, Tree Fl. Malaya 2 (1973) 257; R.J.F.Hend., Fl. Australia 8 (1982) 1; Corner, Wayside Trees Malaya ed. 3, 1 (1988) 349; S.A.Mori & Prance, Fl. Neotrop. Monogr. 21, 2 (1990) 1; Chantar., Kew Bull. 50 (1995) 677; Pinard, Tree Fl. Sabah & Sarawak 4 (2002) 101; H.N.Qin & Prance, Fl. China 13 (2007) 293; Prance in Kiew et al., Fl. Penins. Malaysia, Ser. 2, 3 (2012) 175. — Myrtaceae tribus Lecythideae (A.Rich.) A.Rich. ex DC., Prodr. 3 (1828) 288. — Myrtaceae subtribus Eulecythideae Benth. & Hook.f., Gen. Pl. 1, 2 (1865) 695, nom. inval. — Type: Lecythis Loefl. Napoleaeonaceae A.Rich. in Bory, Dict. Class. Hist. Nat. 11 (1827) 432. — Lecythi- daceae subfam. Napoleonoideae Nied. in Engl. & Prantl., Nat. Pflanzenfam. 3, 7 (1893) 33. — Type: Napoleonaea P.Beauv. Scytopetalaceae Engl. in Engl. & Prantl, Nat. Pflanzenfam., Nachtr. 1 (1897) 242. — Lecythidaceae subfam. Scytopetaloideae (Engl.) O.Appel, Bot. J. Linn. Soc. 121 (1996) 225. — Type: Scytopetalum Pierre ex Engl. Lecythidaceae subfam. Foetidioideae Nied. in Engl. & Prantl, Nat Pflanzenfam. 3, 7 (1892) 29. — Foetidiaceae (Nied.) Airy Shaw in Willis & Airy Shaw, Dict. Fl. Pl., ed. 8 (1973) 465. — Type: Foetidia Comm. ex Lam. For Barringtoniaceae see under subfam. Barringtonioideae. Small to large trees, rarely shrubs. Leaves alternate and distichous or spiral (to rarely opposite and clustered), simple, the margins usually entire, pinnately nerved; stipules absent or minute and caducous. Inflorescences terminal, axillary or cauline, simple racemes, panicles with 2–3 orders of racemose or spicate branches or fascicles, or flowers solitary. Flowers actinomorphic or zygomorphic, hermaphrodite; sepals 2–6 or calyx rarely unlobed; petals 3–6(–8), infrequently 12 or 18, (absent in Foetidia); stamens numerous 10–1210, connate at base into a short or long staminal ring, the ring actinomorphic or prolonged on one side into a strap-like structure arching over the summit of the ovary in some Neotropical species; anthers bilocular, latrorse, introrse or rarely poricidal; ovary inferior or semi-inferior, usually 2–6-locular with 2–115 anatropous ovules in each locule; the axillary placenta at the apex, base or throughout the length of the locule; ovules bitegmic, tenuinucellate; style short or long, undivided. Fruits indehiscent, then dry, fleshy or woody, or dehiscent by a circumscissle operculum (Neotropics), then woody; seeds winged or without wings; endosperm lacking or very scanty; embryo undifferentiated or with plano-convex or foliaceous cotyledons. 1) With contributions by P. Baas & F. Lens (wood anatomy) and R.W.J.M. van der Ham (pollen morphology). — Drawings by M. Anwar, R. van Crevel, Damhuri, A. Fahruroji, H. Jelles, J. Pao, Priyono, Sukirno, R. Wise. Photos on page 116–118 were published with permission from M. Leti, S. Hul, J.-G. Fouché, S.K. Cheng & B. David. 2013. Flore Photographique du Cambodge: 324–326. Editions Privat. 2 Flora Malesiana, Ser. I, Vol . 21 (2013) DISTRIBUTION Twenty four genera in three subfamilies with about 285 species. Only members of the subfamily Barringtonioideae in five genera with 71 species occur in Malesia. The Neotropical species Bertholletia excelsa, Couroupita guianensis, and a few species of Gustavia L. are sometimes cultivated in the region. HABITAT AND ECOLOGY Predominantly a lowland rainforest family ranging from swampy coastal areas to sub- montane forest. Many species are found beside streams and behind mangrove forest. A few high altitude species occur in New Guinea, for example, Barringtonia acutangula subsp. acutangula up to 1600 m, B. monticola up to 2000 m and B. jebbiana at 1450 m. POLLINATION AND DISPERSAL Most species of Barringtonia are night-flowering and visits by bats and moths have been recorded for several species. Norio (2004) recorded the pollination of Barringtonia racemosa by moths in Iriomoto Island, Japan. He concluded that this species was able to establish itself so far out of the range of bats because of the visits from the moths Asota heliconia riukiuana, Erasmia pulchella fritzei, Milionia basalis pyreri and Nevrina procopia. The fruits of some of the lowland species are distributed by water including the two most widespread ones, Barringtonia asiatica and B. racemosa, that have both reached many islands in the Pacific and to Madagascar and in the case of the latter to the East African coast. Stocker & Irvine (1983) reported finding seeds of Barringtonia calyptrata in casowarie dung in Northern Australia and it is probable that some of the New Guinea species are dispersed by these giant birds. References: Norio, T., Pollination of Barringtonia racemosa (Lecythidaceae) by moths on Iromote Island Japan. Ann. Tsukuba Bot. Gard. 23 (2004) 17–20. — Stocker, G.C. & A.K. Irvine, Seed dispersal by cassowaries (Casuarius casuarius) in North Queensland’s rainforests. Biotropica 15 (1983) 170–176. TAXONOMY Traditionally placed in the Myrtales. Thorne (1968) suggested a Thealean relationship and Cronquist (1981) and Takhtajan (1997) both established the order Lecythidales somewhere between the Theales and Malvales. Tsou (1994) was the first to suggest a relationship with the small African family Scytopetalaceae. The Lecythidaceae, Scyto- petalaceae and Napoleonaceae are all closely related and are considered as one family by some authors and separate but closely related families by others. Molecular studies (Anderberg et al. 2002, Schönenberger et al. 2005) have shown the three subfamilies Lecythidoideae, Planchonioideae and Foetidoideae to be a monophyletic group and sister to the Sapotaceae within the broadly defined Ericales sensu APG (1998, 2009). The most recent molecular study of Lecythidaceae is Mori et al. (2007) and it shows the Napoleonaeaceae, Scytopetalaceae and Lecythidaceae conform to highly supported clades. Prance & Kartawinata — Lecythidaceae 3 The Malesian species have generally been referred to the subfamily Planchonioideae of Lecythidaceae, but Reveal (1996) showed that there was an older subfamily name Barringtonioideae, which must now be used and is applied here. The Barringtonioideae has syntricolpate pollen, the cortical bundles have a reversed orientation (with the xylem outside the phloem), the secondary xylem lacks chambered crystals. The Lecythidoideae of the New World has tricolpate pollen, the cortical bun- dles are of normal orientation and the xylem usually has crystal chains of two types. Most of the neotropical genera, including Couroupita and Bertholletia, differ from the Barringtonioideae in the highly zygomorphic flowers with the androecium consisting of a basal ring of stamens and a hood with either more stamens or nectariferous stami- nodes. Gustavia has 6–12 petals whereas the Barringtonioideae has 3–4. The genus Barringtonia was revised by Payens (1967) and recently by Prance (2013) and Planchonia by Kuswata (1965). The Malaysian genus Abdulmajidia was transferred to Barringtonia by Prance (2010), because the only difference establishing the genus was that the fruits had more than one seed. However, some species in Barringtonia have also been found with two or more seeds. References: Anderberg, A.A., C. Rydin & M. Källersjö, Phylogenetic relationships in the order Ericales s.l.: analyses of molecular data from five genes from the plastid and mitochondrial genomes. Amer. J. Bot. 89 (2002) 677–687. — Angiosperm Phylogeny Group (APG), An ordinal classification for the families of flowering plants. Ann. Missouri Bot. Gard. 85 (1998) 531–533. — Angiosperm Phy- logeny Group III, An update of the Angiosperm Phylogeny Group classification for the orders and fami- lies of flowering plants. Bot. J. Linn. Soc. 161 (2009) 105–121. — Cronquist, A., An integral system of classification of flowering plants. Columbia Univ. Press, New York (1981). — Kuswata Kartawinata, E., The genus Planchonia Blume (Lecythidaceae). Bull. Bot. Surv. India 7 (1965) 162–187. — Mori, S., C.-H. Tsou, B. Cronholm & A.A. Anderberg, Evolution of Lecythidaceae: information from combined ndhF and trnL-F sequence data. Amer. J. Bot. 94 (2007) 289–301. — Payens, J.P.D.W., A monograph of the genus Barringtonia (Lecythidaceae). Blumea 15 (1967) 157–263. — Prance, G.T., Notes on the Lecythidaceae of Peninsular Malaysia. Blumea 55 (2010) 14–17. — Prance, G.T., A revision of Barringtonia (Lecythidaceae). Allertonia 12 (2013) 1–184. — Reveal, J.L., Latest news on vascular plant family nomenclature. http://www.plantsystematics.org/reveal/pbio/fam/OLDNEWS98.html. 1996, last updated 1 Jan. 1999, last checked May 2012. — Schönenberger, J., A.A. Anderberg & K.J. Systma, Molecular phylogenetics and patterns of floral evolution in the Ericales. Int. J. Pl. Sci. 166 (2005) 265–288. — Takhtajan, A., Diversity and classification of flowering plants. Columbia Univ. Press, New York (1997) — Thorne, R.F., Synopsis of a putative phylogenetic classification of flow- ering plants. Aliso 6 (1968) 57–66. — Tsou, C.-H., The embryology, reproductive morphology, and systematics of Lecythidaceae. Mem. New York Bot. Gard. 71 (1994) 1–110. REPRODUCTIVE MORPHOLOGY The embryology and reproductive morphology of the entire Lecythidaceae
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