Bertholletia, Bonpland, 1808

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 : 91-95

Bertholletia clade (63% BS; Figs. 2 , 4–15 )

Our results support the monophyly of the Bertholletia clade, which includes Bertholletia , Corythophora , Eschweilera and Lecythis as defined in Mori & Prance (1990), and the results are congruent with Mori et al. (2007) and Huang et al. (2011). The Bertholletia clade has a distinctive combination of its own synapomorphies and character states shared with some of the outgroup taxa. Character optimization using the unambiguous option of Winclada shows that morphological synapomorphies supporting the monophyly of the clade include the presence of a two or four-locular ovaries (character 39), the presence of an aril (character 47), and the absence of cotyledons (character 49). In addition, flowers of the Bertholletia clade have a distinctive combination of androecium features shared with some outgroup taxa, including zygomorphy (character 20), presence of a hood (= coiled ligule, character 26), the presence of stamens in the hood (character 32; called “vestigial” stamens because they lack anthers), and the absence of an external flap (character 25). None of the outgroup taxa has this combination of characters; all are either actinomorphic or all of the appendages on the hood are either staminodes (species of Couroupita ) or vestigial stamen nectaries as in Couratari . In addition, there are no species in the Bertholletia clade with staminal tubes, as in species of Allantoma , Cariniana , and Gustavia (character 21).

There are also seed features that separate the Bertholletia clade from the outgroup. For example, there are no species in the Bertholletia clade with linear seeds with a notch at the base (cf. Allantoma lineata [Mart. ex O. Berg [1858: 508]] Miers [1874: 297]), and none have fleshy cotyledons (cf. Gustavia ; character 49), winged seeds (unilaterally winged in Allantoma and Cariniana , circumferentially winged in Couratari ; character 45), seeds with long trichomes extending from the seed coat (cf. Couroupita ; character 44) ( Tsou & Mori, 2002), or seeds with leaf-like cotyledons (cf. Cariniana , Couratari , and Couroupita ; character 49). The only genus outside of the Bertholletia clade with a ligule extending from a staminal ring, a feature common to all members of the Bertholletia clade, is Couroupita (character 24). Within the Bertholletia clade, the four genera are divided into the ten clades described and illustrated below ( Figs. 4–14 ).

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 4. The Lecythis pisonis clade (see Fig. 101 in Mori & Prance, 1990 for vouchers). A. Flower with an open androecial hood. B. Medial longitudinal section of flower with flat androecial hood. The androecial hood possesses a proximal group of staminodes for about one-quarter of its length (not distinguishable in this image) and vestigial stamens for the rest of the length. C. Medial longitudinal section of ovary. Note the stylar collar located just under the stigma. D. Cross-section of 4-locular ovary. E. Operculum with a woody columella that projects into the fruit. F. Fruit base. The fruits of this clade are the largest of all Lecythidaceae. G. Sulcate seed with a long basal aril. Note the funicle projecting from the bottom of the aril. Drawing by B. Angell.

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 10. The Corythophora clade. A–E. C. amapaensis (see Fig. 42 in Mori & Prance, 1990 for vouchers), F–L. C. alta (see Fig. 12 in Mori & Lepsch-Cunha, 1995 for vouchers) A. Apical view of flower. Note the closed androecial hood typical of all species in this clade. B. Inferior ovary with all but four calyx-lobes removed. Note that the calyx-lobes are strongly imbricate. C. Cross-section of 2-locular ovary. D. Medial section of an androecium with the androecial hood dorsiventrally thickened. E. Campanulate fruit base. F.Apical view of flower. G. Ovary, calyx, and style of a very young fruit. Note that the calyx-lobes are not imbricate. H. Cross-section of 2-locular ovary. I. Medial longitudinal section of androecium with dorsiventrally thickened hood. J. Cylindrical fruit. K. Operculum without a columella. L. Seeds with basal arils. The seeds of this clade are oblong to fusiform. Drawings A–E by B. Angell and F–L by A. Tangerini.

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 12. Arils found in the Eschweilera integrifolia clade. A. E. caudiculata (Cornejo 8106). B. E. antioquensis (Luteyn & Callejas 12025). C. E. ovalifolia (S. A. Mori photo only). A. Top left: Side view of seed showing lateral aril. Note that this type of lateral aril extends onto both ends of the seed. Top right: Rounded side of hemispherically-shaped seed. Bottom left: Lateral aril of wedge-shaped seed. Bottom right: Flat side (left) and round side (right) of hemispherically shaped seed. B. Three different views of wedge-shaped seed. C. Cross-section of seed with spreading aril. Note that the next layer is the seed coat, and the center is filled with an undifferentiated (= macropodial) embryo. Photos by S. A. Mori.

FIGURE 13. The E. tetrapetala clade.A–B. E. tetrapetala (see Fig. 63 in Mori & Prance, 1990 for vouchers), C. E. alvimii (see Fig. 65 in Mori & Prance, 1990 for vouchers). D–E. E. nana (see Fig.8 in Prance & Mori, 1991 for vouchers).A. Medial section of androecium. Note that the androecial hood has a single coil and that there are vestigial stamens on both the exterior and interior of the coil, a feature unique to all species of this clade. B. Cross-section of 2-locular ovary, a feature common to all species of this clade. C. Medial section of androecium of E. alvimii. D. Fruit base and operculum. Note that the operculum lacks a columella. E. Seeds with basal arils. Drawings by B. Angell.

FIGURE 14. The Lecythis corrugata clade.A. L. corrugata subsp. corrugata (Mori et al. 25730). B. L. confertiflora (Mori et al. 20801). C. L. persistens subsp. persistens (Mori et al. 25651). D. L. persistens subsp. aurantiaca (Mori et al. 24724). E. Lecythis idatimon (Mori et al. 25745). F. L. persistens subsp. persistens. A. Apical view of flower. Note that this is the only species in this clade with a closed androecial hood and without lateral flanges. B. Anterior view of flower. Note that this species has an open androecium and the sides of the ligule with lateral flanges. C.Anterior view of flower. Note that his species has an open androecium and the sides of the ligule possess lateral flanges. D. Lateral view of flowers. Note that this species has an open androecium and the sides of the ligule with lateral flanges. E. Young fruits with tuberculate pedicels and hypanthia. F. Young fruits with rugose pericarp and fusiform seeds with a small aril at their base. Photos by S. A. Mori (A, C–F) and C. Gracie (B).

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..