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A Morphological Cladistic Analysis of Lecythidoideae with Emphasis on Bertholletia, Corythophora, Eschweilera, and Lecythis

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A Morphological Cladistic Analysis of Lecythidoideae with Emphasis on Bertholletia, Corythophora, Eschweilera, and Lecythis A morphological cladistic analysis of Lecythidoideae with emphasis on Bertholletia, Corythophora, Eschweilera, and Lecythis 1 2 3 YA-YI HUANG ,SCOTT A. MORI , AND LAWRENCE M. KELLY 1 The New York Botanical Garden, 2900 Southern Blvd., Bronx, NY 10458-5126, U.S.A. Current address: Biodiversity Research Center, Academia Sinica, Taipei 11529, Taiwan; e-mail: lecy.yhuang@ gmail.com 2 Institute of Systematic Botany, The New YorkBotanical Garden, 2900 Southern Blvd., Bronx( NY 10458-5126, U.S.A.; e-mail: [email protected] 3 The New York Botanical Garden, 2900 Southern Blvd., Bronx( NY 10458-5126, U.S.A.; e-mail: [email protected] Abstract. A cladistic analysis was conducted to test the monophyly of Eschweilera and Lecythis as well as to examine the relationships of these two genera and their close relatives Bertholletia and Corythophora. The study included 86 species, represent- ing all four genera and covering the range of taxonomic and morphological variation in the genera. The data matrix included 49 parsimony-informative characters derived from vegetative, floral, fruit, and seed morphology and anatomy. The results based on the consensus of all most parsimonious trees indicate that Bertholletia, Corythophora, Eschweilera,andLecythis form a clade supported by brachyparacytic stomata, the absence of pedicels (with subsequent reversals in several clades), a two or four- locular ovary, the presence of an aril, and the absence of cotyledons. Within the clade, the monophyly of Corythophora is supported by the presence of inflores- cence scales and the absence of nectar. Eschweilera is monophyletic only if E. congestiflora and E. simiorum are excluded. The monophyly of Eschweilera is supported by the presence of a two-locular ovary. Lecythis is not monophyletic, but sections Corrugata, Pisonis,andPoiteaui are monophyletic. Three species of section Lecythis aremorecloselyrelatedtoEschweilera, and other species of section Lecythis along with Bertholletia excelsa remain as unresolved. Key Words: Bertholletia, Corythophora, Eschweilera, Lecythis, Lecythidoideae, zygomorphy. Lecythidaceae are small to large trees that &Mori,1979; Mori & Prance, 1990b;Moriet grow in lowland rainforest (Mori & Prance, al., 2007; Huang et al., 2008). Species of 1990b). The family consists of three subfa- the Lecythidoideae are confined to tropical milies (Mori et al., 2007): Foetidioideae, America. The Lecythidaceae is monophyletic Planchonioideae, and Lecythidoideae. The and each of the subfamilies is monophyletic Foetidioideae consists of 17 species in one (Morton et al., 1998;Morietal.,2007; APG III, genus. These species are found in East Africa, 2009). The relationships within the subfamilies, Madagascar, and Mauritius (Mori et al., however, are not fully understood, especially 2007). The Planchonioideae consists of 59 within the Lecythidoideae. species in six genera. They are found in tropical Lecythidoideae includes both endemics with Asia, tropical Africa, Australia, and the Pacific narrow distributions and widespread taxa. They Islands (Mori et al., 2007). The Lecythidoideae occupy diverse habitats but are most abundant consists of ca. 210 species in 10 genera (Prance in non-flooded primary forests (Mori & Prance, Brittonia, 63(3), 2011, pp. 396–417 ISSUED: 1 September 2011 © 2011, by The New York Botanical Garden Press, Bronx, NY 10458-5126 U.S.A. 2011] HUANG ET AL.: MORPHOLOGICAL ANALYSIS OF LECYTHIDOIDEAE 397 1990b). Species of Lecythidoideae are charac- However, the type species of Eschweilera and terized by fibrous bark; simple, alternate leaves; Lecythis were not included and some critical actinomorphic or zygomorphic androecia; taxa were not sampled, e.g., Eschweilera numerous stamens; inferior or half-inferior section Te t r a p e t a l a S. A. Mori. In this analysis, ovaries; bitegmic ovules; and the presence of Bertholletia, Corythophora, Eschweilera,and cortical bundles (Prance & Mori, 1979). Lecythis together were monophyletic (the The taxonomy of Lecythidoideae places great Bertholletia clade), but the phylogenetic rela- emphasis on floral features, especially on the tionships among them were not fully resolved structure of the androecium. The ten genera of (Fig. 1 in Mori et al., 2007). Within the Lecythidoideae can be divided into two groups Bertholletia clade, moreover, the results further based on androecial symmetry. Three genera indicated that Eschweilera and Lecythis may have actinomorphic androecia: Allantoma not be monophyletic. Miers (eight species; Huang et al., 2008), Grias Of the four genera, Lecythis was described L. (nine species; Mori 1979b; Clark & Mori, first. In 1758, two years after Loefling died, 2000; Cornejo & Mori, 2010;Morietal.,2010), Linnaeus published Loefling’s description of and Gustavia L. (41 species in three sections; a new genus, Lecythis Loefl., but the first Mori, 1979a;Morietal.,2007). Seven genera species in the genus (Lecythis ollaria L.) was have zygomorphic androecia: Bertholletia not published until 1759 (Dorr & Wiersema, Bonpl. (one species; Prance, 1990b), Cariniana 2010). These were the first valid publica- Casar. (nine species; Huang et al., 2008), tions of Lecythidaceae, even before the Corythophora R. Knuth (four species; Mori & family status was established. Lecythis is Prance, 1990c), Couratari Aubl. (19 species in the third largest genus of Lecythidoideae; three sections; Prance, 1990c), Couroupita however, it is the most diverse of all genera Aubl. (three species; Prance, 1990a), Eschwei- in terms of floral and fruit features (Mori, lera Mart. ex DC. (93 species in four sections; 1990c). Species of Lecythis are distributed Mori & Prance, 1990d;Mori,1992, 1995a, from Nicaragua to Rio de Janeiro, Brazil 2007), and Lecythis Loefl. (27 species in four (Fig. 27 in Mori & Prance, 1990b) sections; Mori, 1990c, 1995b). Based on Bertholletia was described by Humboldt and comparative morphology, Mori et al. (2007) Bonpland (1807). Bertholletia excelsa Bonpl. hypothesized that the actinomorphic androe- (the Brazil nut) is the only species of the genus. cium is the plesiomorphic state in the Lecythi- It is also the best known species in the family doideae. Most actinomorphic-flowered genera because of its edible seeds (Prance, 1990b). have their greatest diversity west of the Andes Bertholletia excelsa can be easily distinguished or in western Amazonia, whereas the zygomor- from the other three genera by two instead of phic-flowered genera are most diverse in six calyx lobes, the operculum falling inside central Amazonia and the Guianas. instead of outside the fruit, and by the absence Morton et al. (1998) conducted a phyloge- instead of the presence of a fleshy aril. It is netic analysis to test the monophyly of Lecy- widespread throughout Amazonia and the thidaceae and the monophyly of each of the Guianas (Fig. 21 in Mori & Prance, 1990b). three subfamilies based on morphological, Knuth (1939) established Corythophora based anatomical, cytological, and plastid rbcL and on Corythophora alta R. Knuth. He separated trnL sequence data. This study included one Corythophora from Lecythis because all the species to represent each genus, and one hood appendages of C. alta possess anthers, individual was sampled for each species. The whereas most species of Lecythis do not have results indicated that Lecythidaceae are mono- hood anthers or only part of the hood appen- phyletic and each of the three subfamilies is monophyletic (Fig. 1 in Morton et al., 1998). dages have anthers (Mori & Prance, 1990c). He Mori et al. (2007) performed another analysis separated Corythophora from Couroupita by its to test the monophyly of genera of the dehiscent versus indehiscent fruits (Mori & subfamily Lecythidoideae based on plastid Prance, 1990c). Species of Corythophora are ndhF and trnL-F sequence data. This study limited to Surinam, French Guiana, and Brazil- incorporated more species into the analysis. ian Amazonia from Manaus to Amapá, where 398 BRITTONIA [VOL 63 they are found mostly north of the Amazon The taxonomy follows Prance and Mori River (Fig. 23 in Mori & Prance, 1990b). (1979) and Mori and Prance (1990b). The Eschweilera was first described by De sampled taxa are listed in Table I. — Candolle (1828). Martius (1837) accepted Data collection. Morphological and ana- Eschweilera as a genus and listed several tomical characters were collected based on fi features of Eschweilera distinct from Lecy- original observations in the eld and the this (e.g., bilocular ovary). However, many images available on the Lecythidaceae Pages subsequent authors still considered (http://sweetgum.nybg.org/lp/index.html). Eschweilera as part of Lecythis (Endlicher, Some characters were derived from the 1840;Berg,1856, 1857, 1858). In 1874, literature, herbarium specimens at INB, INPA, Miers maintained the generic status of MO, NY, and US, or pickled collections at Eschweilera and provided a list of features NY. that distinguished Eschweilera from Lecy- Leaf fragments used for study of venation this. The generic status of Eschweilera has patterns and stomata were placed in vials not changed since then. Eschweilera is the with 50% alcohol and bathed in boiling largest and the most widely distributed water for ten minutes. This procedure was genus of Lecythidoideae. Species of repeated two or three times until no further Eschweilera range from Veracruz, Mexico, color leached into the solution. After cool- to Rio de Janeiro, Brazil (Fig. 25 in Mori & ing to room temperature, samples were Prance, 1990b). washed with water two or three times, and In the present
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