Eschweilera parvifolia, Mart. ex Candolle, 1828

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 : 111

Eschweilera parvifolia clade (94% BS; Fig. 2B , Fig. 15 )

This clade consists of a sample of 29 of the approximately 63 species (minus the species now considered as belonging to the E. integrifolia clade) recognized by Mori & Prance (1990). Species of this clade are found nearly everywhere in the Neotropics, ranging from Veracruz, Mexico to Rio de Janeiro, Brazil ( Huang, 2010).

The sections of Eschweilera , as defined by Mori & Prance (1990), include Eschweilera sect. Tetrapetala (our E. tetrapetala clade discussed above); Eschweilera sect. Jugastrum , consisting only of E. tenuifolia (O. Berg 1858: 502) Miers (1874: 266) ; Eschweilera sect. Bracteosa , consisting of the sampled E. bracteosa (Poepp ex. O. Berg (1856: 455) Miers (1874: 274), E. laevicarpa S. A. Mori (1987: 32) , and E. cyathiformis S. A. Mori (1989: 20) , and the nonsampled E. rabeliana S. A. Mori (1989: 21) and E. revoluta S. A. Mori (in Mori & Prance 1990: 174); and Eschweilera section Eschweilera with the remaining species (minus those found in the E. integrifolia clade). The type, E. parvifolia ( Mori & Prance, 1990) , is found in this clade.

Mori & Prance (1990 d) included species of our E. integrifolia (described above), E. tetrapetala (described above), and the E. parvifolia clades in their concept of Eschweilera . Based on our results, Eschweilera is not monophyletic.

The most useful morphological synapomorphy of the E. parvifolia clade is the presence of a lateral aril (character 48, Fig. 15F ). Although there are a few species with lateral arils in the E. integrifolia clade, most of those species have spreading arils ( Fig. 12C ) and the ones with lateral arils are either much larger and/or wrap around the ends of the seeds ( Figs. 12A, 12B , see above discussion of the E. integrifolia clade). In addition, this is the only clade with consistently double-coiled androecial hoods ( Fig. 15B–H ) in contrast to the consistently single-coiled androecial hoods of the Eschweilera tetrapetala and the triple-coiled androecial hoods of the E. integrifolia clades. The species of the Eschweilera parvifolia and E. integrifolia clades are the only species to have vestigial stamen nectaries in the Bertholletia clade, a feature that is also found outside of the clade in Couratari (Mori et. al., 2015).

Eschweilera tenuifolia differs from the other species of this clade because it lacks pedicels (character 13) and its seeds do not have an aril (character 47). Other defining features are wedge-shaped seeds; a corky seed coat; and seed germination from the sides (Fig. 18M in Prance & Mori, 1979) instead of the ends of the seeds (not coded). The flowers of E. tenuifolia have a double-coiled androecial hood identical to the other species of the E. parvifolia clade. This species was the only one included in E. section Jugastrum by Mori & Prance (1990) and can only be recognized as a separate section if the poorly understood basal clade (the E. mexicana clade) is also recognized as a section but that clade has no differences from the other species of the Eschweilera parvifolia clade. In this study, the representatives of Eschweilera section Bracteosa fall into two unresolved clades. Whether this section should be recognized is open to question, but it is unlikely that the persistent bracts and bracteoles in the inflorescences are stable enough for use in defining taxonomic groups ( Fig. 2B ).

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 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 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..