Lecythis chartacea, O. Berg, 1856

Huang, Ya-Yi, Mori, Scott A. & Kelly, Lawrence M., 2015, Toward a phylogenetic-based Generic Classification of Neotropical Lecythidaceae- I. Status of Bertholletia, Corythophora, Eschweilera and Lecythis, Phytotaxa 203 (2), pp. 85-121 : 102-104

Lecythis chartacea clade (76% BS; Figs. 2A , 8 )

This clade is distributed in Amazonian Venezuela, the Guianas, and in western to eastern Amazonian Brazil ( Huang, 2010). None of the morphological characters that were included in the analysis provide synapomorphies for this clade, and the only apparent morphological distinction for the clade is the more-or-less fusiform seeds with salient longitudinally oriented major veins and the areas between them with salient higher order veins ( Figs. 8F ). These seeds differ from the smooth inter-venal areas of the seeds of the L. ollaria clade ( Fig. 9A–C ), the dendritically arranged pattern and plane or impressed veins of the L. poiteaui clade ( Fig. 9M–P ), and the hard seed coat of the Bertholletia excelsa clade ( Fig. 7G ). Members of the L. chartacea clade possess an androecial hood with swept in appendages ( Figs. 8A, I, L ) as do some of the species of the L. poiteaui ( Fig. 6D ) and B. excelsa ( Fig. 7D ) clades. The hood of the L. ollaria clade differs from these clades in its possession of a single coil ( Fig. 5A, G ). In addition, zygomorphic-flowered species with these types of androecial hoods do not possess obvious vestigial stamen nectaries, like those of the Eschweilera integrifolia ( Figs. 11B, F ) and E. parvifolia ( Figs 15B, H ) clades and the outgroup genus Couratari . The presence of mucilage ducts in the ovary and/or the calyx lobes (character 17) is found in both the L. poiteaui ( Figs. 6E, H, L ) and L. chartacea clades but they are more common in the former clade; relatively long, obliquely oriented or geniculate styles occur in the L. poiteaui ( Fig. 6G ), B. excelsa ( Fig. 7B ), and L. chartacea ( Figs. 8B, 8J ) clades; indehiscent fruits are found in some of the species of the L. poiteaui , B. excelsa , and some of the species of the L. chartacea clades. Moreover, there are both dehiscent- and indehiscent-fruited species in the L. poiteaui and L. chartacea clades. In these clades, the fruits are of two types: they can be large with a relatively thin pericarp and fall to the ground without dehiscing (e.g., L. lurida and L. prancei Mori [1990b: 304] , Fig. 6I ) or the fruits dehisce but do not release the seeds, which are so large that they do not fall from the fruit (e.g., L. ibiriba ( Miers 1874: 236) Smith et al. [2013: 447] , Fig. 6N ). In the Lecythis poiteaui clade, regardless of fruit type (whether truly indehiscent or with seeds that remain stuck inside the fruit), the seeds are large, more-or-less round (i.e., not markedly longer than broad), have plane or slightly impressed, dendritic veins, and a vestigial ( Fig. 6P ) aril or no sign of an aril ( Figs. 6J , 9M–P ).

Indehiscent fruits of the L. chartacea clade are possessed by the riverine species L. rorida (mistakenly treated as a synonym of L. chartacea by Mori, 1990b), which has fruits that usually drop into the water with the non-arillate seeds trapped inside, and the terra firme species L. gracieana S. A. Mori (in Mori & Lepsch-Cunha 1995: 47) and L. parvifructa S. A. Mori (1990b: 312) , which have relatively small, single-seeded fruits that fall to the ground at maturity without dehiscing. All of the remaining species sampled in the L. chartacea clade have dehiscent fruits and seeds with well-developed basal arils.

Lecythis brancoensis is sister to all other species of the L. chartacea clade ( Fig. 2A ), but was included in Lecythis Section Poiteaui by Mori (1990b). It differs from other species of the L. chartacea clade in the presence of anthers or antherodes (character 32) on the innermost appendages of the androecial hood and the absence of a closed androecium (character 33). It was placed in Lecythis sect. Poiteaui based on the hypothesis that L. brancoensis is also bat-pollinated, which is supported by its unbranched terminal inflorescence and very large numbers of stamens, In addition, L. brancoensis shares a papillate abaxial leaf surface with the bat-pollinated L. barnebyi and L. poiteaui . In Huang et al. (2011), L. brancoensis was recovered as a clade with the two known bat-pollinated species of Lecythis sect. Poiteaui ; however, the current study does not support the relationship between the bat-pollinated species of the Lecythis poiteaui clade ( Fig. 2A ) and the hypothetical bat-pollinated L. brancoensis of the L. chartacea clade.

Eschweilera congestiflora and E. simiorum were placed in Eschweilera Section Eschweilera by Mori & Prance (1990) but these two species possess features that are common for species of the L. chartacea clade, e.g., a non-coiled ligule (character 26; Fig. 8C, I, L ), curved inward appendages arising from the apex of the ligule (character 31), a 4- locular ovary (character 39), and seeds with a basal aril (characters 47, 48; Fig. 9L ). Mori et al. (2007) pointed out that these two species were placed in the wrong genus as indicated by molecular data. In this study, these two species are embedded in the L. chartacea clade, but new combinations will not be needed because they were originally described as L. congestiflora Benoist (1915: 177) and L. simiorum Benoist (1915: 178) ( Fig. 2 in Mori et al., 2007).

have more-or-less globose seeds, plane or impressed veins, and the overall dendritic venation pattern. Photos by S. A. Mori.

Figures

FIGURE 2A. Strict consensus of 66 most parsimonious (MP) trees based on total evidence. Bootstrap values (>50%) are given above the branches.All clades in this figure are part of the Bertholletia clade. The Lecythis pisonis, L. ollaria, Bertholletia excelsa, L. poiteaui, and Corythophora clades of the Bertholletia clade are shown.

FIGURE 5. The Lecythis ollaria clade. A–F. L. minor (A based on Galdames 5768; B–F. based on Nee & Mori 3580). G–J. L. tuyrana (based on Galdames 5766). A. Medial longitudinal section of flower showing the single coil and the vestigial stamens limited to the exterior of the coil. B. Apical view of dehisced fruit. C. Lateral view of fruit. D. Basal view of operculum. E. Apical view of operculum. F. Seeds with basal arils. G. Medial longitudinal section of flower showing single but short coil. H. Apical view of fruit with operculum removed and showing seeds inside. I. Basal view of operculum. Note that the columella is not developed. J. Seed with basal aril (above). Note funicle protruding from aril. Photos A and G–J by C. Galdames and photos B–F by S. A. Mori.

FIGURE 6. The Lecythis poiteaui clade. A–C. L. barnebyi (A–B see Mori & Lepsch-Cunha, 1995 for vouchers and C see Mori 1990c for voucher). D–E. L. prancei (see Mori, 1990c for vouchers). F–J. L. lurida (see Mori, 1990c for vouchers). K–L. L. ibiriba (K–L based on Popovkin 496, M based on Popovkin 497, N–P based on Cardoso 2338). A. Anterior view of flower showing open androecial hood. B. Lateral view of flower showing open androecial hood. C. Medial section of androecium. Note that innermost hood appendages arise from slight expansion of hood, there is a long anterior hood extension, and all of the appendages are swept inward. D. Medial section of androecium. Note that the appendages are swept inward. E. Cross-section of ovary showing mucilage ducts in the ovary wall. F. Medial section of flower. Note that hood appendages are swept inward. G. Medial section of calyx and ovary. Note that style is obliquely oriented and long. H. Cross-section of ovary. Note mucilage ducts in the calyx-lobes. I. Indehiscent fruit, operculum facing downward. J. Seed with dendritic venation and without well-developed aril. K. Medial section of flower. Note that hood appendages are swept inward. L. Cross-section of ovary showing mucilage ducts in ovary wall. M. Lateral view of fruit. N.Apical view of open fruit with seeds inside. O. Basal view of operculum. P. Seed with dendritic venation and poorly-developed aril. Drawings A–B by A. Tangerini and all others by B. Angell.

FIGURE 7. The Bertholletia excelsa clade (see Fig. 45 in Mori & Prance, 1990 for vouchers except for B which is vouchered by Mori et al. 17503). A. Flower showing petals tightly pressed against androecium and turned downward at their apices. B. Calyx, ovary, and style. Note that the calyx consists of two lobes, the ovary is inferior and very short and the style is oblique. C. Cross-section of 4-locular ovary. D. Medial section of androecium showing swept in vestigial stamens and the anterior ligular extension. E. Fruit showing that the opercular opening is smaller in diameter than the diameter of the seeds. F. Operculum. Note that it drops into the inside of the fruit a maturity. G. Seed. This is the only neotropical Lecythidaceae with a ligneous seed coat. H. A seedling. The embryo lacks cotyledons and is mostly composed

FIGURE 8. The Lecythis chartacea clade.A–E. L. chartacea (A–C based on Mori et al. 26485, D–F based on Nee & Mori 4199). G–K. L. gracieana (see Fig. 19 in Mori & Lepsch-Cunha, 1995 for vouchers). L. L. holcogyne (see Fig. IV-1 in Mori, 1987 for voucher). A. Medial section of flower. Note that the vestigial stamens arise from the apex of the ligule and are swept inwards. B. Lateral view of ovary with all other floral parts removed. Note the geniculate style. C. Cross-section of 4-locular ovary. D. Lateral view of fruit. E. Operculum. F. Two seeds. The salient major veins run parallel to the main axis of the seed and the secondary veins depart from them into the area between the major veins. G.Apical view of flower. H. Lateral view of flower showing petals tightly pressed against androecium making it difficult for all but robust bees to enter the flower. I. Medial section of an androecium showing vestigial stamens swept inward. J. Medial section of ovary. Note obliquely oriented style. K. Cross-section of 4-locular ovary. L. Medial section of androecium showing swept inward vestigial stamens (a) and anterior hood extension (b). Drawings A–C by C. Carollo, the remaining by B. Angell, and the photographs by S. A. Mori.

FIGURE 9. Seeds of the Lecythis ollaria (A–C), L. chartacea (D–L), and Lecythis poiteaui (M–P) clades. Lecythis ollaria clade—A. L. ollaria (Davidse & González 12096A). B. L. minor (Prance 23172). C. L. tuyrana (not vouchered). All species in this clade have a tendency toward more-or-less fusiform seeds, plane primary veins, major veins oriented along length of seeds, and higher order veins absent, i.e., the areas between the major veins are smooth. Most of the veins of L. ollaria and L. minor only extend for part of the length of the seeds while those of L. tuyrana extend from the base to the apex of the seed. Lecythis chartacea clade—D. L. chartacea (Nee & Mori 4199). E. L. alutacea (Redden et al. 1732). F. L. brancoensis (Silva 48). G. Lecythis rorida (Mori et al. 20428). H. L. retusa (Ramos s.n. NY barcode 00684323). I. L. gracieana (Freitas et al. 745). J. Lecythis parvifructa (Freitas et al. 726). K. L. holcogyne (Mori & Pipoly 15493). L. Eschweilera. simiorum (Clark 4333). Most species in this clade have fusiform or, less frequently, globose seeds, salient primary veins, major veins oriented along the entire length of the seeds, and salient higher order veins. The seeds associated with indehiscent fruits (e.g., those of L. gracieana (I) and L. parvifructa (J), tend to be more globose. Lecythis poiteaui clade—M. L. barnebyi (Costich & dos Santos 836). N. Lecythis ibiriba (Carvalho et al. 6026). O. L. lurida (Prance 26574). P. L. prancei (Mori 20286). All species in this clade

FIGURE 11. The Eschweilera integrifolia clade. A–D. E. integrifolia (based on Cornejo 8111). E–H. E. jacquelyniae (E–F based on Hernández 828 and G–H based on Galdames 6142). I–J. E. ovalifolia (based on an unvouchered photo by S. A. Mori). A. Apical view of flower. Note the closed androecial hood typical of all species of this clade. B. Medial section of triple-coiled androecial hood. Note that there are three coils, a feature common to this clade, and that the coils are oriented horizontally. C. Lateral view of fruit. D. Apical view of a fruit with “a” marking a seed completely surrounded by the aril (= spreading aril). E. Cauline inflorescences. Note that this inflorescence is very near to the ground but other inflorescences of this species may also occur on the branches. F. Medial section of triple coiled androecium. Note that the coils are oriented vertically. G.Apical view of opened fruit with “a” marking a seed with a spreading aril. The aril is pale yellow and the immature seeds are white. H. Inside view of operculum. I. Apical view of open fruit. Note that the seeds are completely surrounded by arils. J. Cross-section of seed. Note the white layer on the outside (= aril), the seed coat, and the solid embryo which does not have differentiated cotyledons. Photos A–D by X. Cornejo, E–H by F. Hernández, and I–J by S. A. Mori.

FIGURE 15. The Eschweilera parvifolia clade.A–G. E. pedicellata (see Fig. 85 in Mori & Prance, 1990 for vouchers). H–J. E. ovata (see Fig. 87 in Mori & Prance for vouchers). K. E. romeu-cardosoi (see Mori & Lepsch-Cunha, 1995 for vouchers).A. Apical view of flower. Note that this and all species of this clade have a closed androecium. B. Medial longitudinal section of an androecium. Note the incipient third coil in contrast to the double coil of most species of this clade. C. Cross-section of 2-locular ovary. Although most species of this clade have 2-locular ovaries there are a few species that are 4-locular. D. Base of fruit. E. Operculum of fruit. F. Lateral view of seed with lateral aril. G. Basal view of seed showing position of lateral aril. H. Medial longitudinal section of androecial hood showing double coil. I. Stamen showing clavate filament. J. Medial longitudinal section of ovary showing basal insertion of the ovules. K. Seedling. Note that the embryo remains inside of the seed coat upon germination. Line drawings by B. Angell..