Eschweilera integrifolia

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 : 106-107

Eschweilera integrifolia clade (<50% BS; Fig. 2B , 11 , 12 )

This clade comprises 19 sampled species of Eschweilera included in Eschweilera section Eschweilera by Mori & Prance (1990). Species of this group are found from central to western Amazonian Brazil, the Andes, forests of the Pacific coasts of Colombia, Ecuador, and Central America as far north as Costa Rica.

This clade is defined by a triple coil (characters 26, 27) with vestigial stamen nectaries at the apex of the last coil (not coded; Fig. 11B, F ). In addition, most of the species (e.g., E. aguilarii S. A. Mori [2007: 903] , E. amplexifolia S. A. Mori [ Mori & Prance 1990: 201], E. andina ( Rusby 1896: 37) Macbride [1941: 246] , E. collinsii Pittier (1908: 97) , E. integrifolia , E. ovalifolia ( Candolle 1828: 292) Niedenzu [1892: 40] , and E. sessilis A. C. Smith 1933: 21 ) have a spreading aril that completely surrounds the seed (character 48; Figs. 11D, G, I , 12C ), but several species (e.g., E. antioquensis Dugand & Daniel [1938: 1] , E. caudiculata R. Knuth [1939: 95] , and E. rimbachii Standley [1935: 31] ) possess arils that are lateral but differ from the lateral arils of the E. parvifolia clade by having their ends extend around the base and apex of the seed ( Fig. 12A, B ); one species ( E. jacquelyniae S. A. Mori [ Mori & Prance 1990: 192]) has very large and fleshy lateral arils ( Fig. 11D ).

Eschweilera amazoniciformis , endemic to central Amazonian Brazil, is sister to the remaining species of the clade ( Fig. 2B ). This species is distinguished by the presence of four instead of six calyx-lobes (character 16) and four instead of six petals (18). In addition, it is the only known species of neotropical Lecythidaceae with the combination of a triple-coiled androecial hood and fusiform seeds with a well-developed basal aril.

Most of the species of the E. integrifolia clade are found in western Amazonia and the mountain valleys and slopes of the Andes, with the exception of the central Amazonian E. amazoniciformis and Eschweilera ovalifolia . The Andean and western Amazonian species possess predominantly red flowers, but some species, for example, the coastal Ecuadorean species E. awaensis S. A. Mori & Cornejo (2011: 470) and the western to central Amazonian species E. ovalifolia , have yellow flowers.

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 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.