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.