Premise of research. Previous studies of secretory structures in species of the Neotropical dragon's blood Croton (section Cyclostigma) show inconsistencies in their classification. An accurate assessment of the identity and homology of such structures is essential for taxonomic and evolutionary studies. Methodology. Field-collected leaves, stipules, and flowers at different developmental stages were sampled. The material was subjected to standard anatomical study by light microscopy and SEM, and secretions were evaluated by histochemical analyses. Pivotal results. Leaves and flowers of Croton echinocarpus and Croton urucurana present five secretory structures (idioblasts, laticifers, colleters, extrafloral nectaries, and floral nectaries) with high similarity between the two species. Idioblasts secrete compounds of a mixed nature; laticifers are branched and nonarticulated; and colleters and nectaries present hydrophilic secretion. The leaf marginal glands previously described as extrafloral nectaries are actually colleters of the standard type. We found colleters in staminate and pistillate flowers. The histochemical tests detected proteins in the secretions of all structures. Conclusions. The classes of secondary metabolites detected support the biological activities of secretion described in the literature. The correct identification of colleters in flowers establishes a new register of these structures in flowers of this genus. We show that an approach integrating anatomic structure, histochemistry, and period of secretion activity allows for a more accurate classification and homology assessment of secretory elements in this genus, which is exceptionally rich in this type of structures.
Establishing species limits in clades in Croton using characters from external morphology and common molecular markers has proved cumbersome, especially in Croton section Cyclostigma, a group of 50 Neotropical arborescent species commonly known as dragon's blood. Given this, we explored leaf and shoot apex anatomical characters for their utility in distinguishing species or groups of species in this section. We analysed 90 specimens belonging to section Cyclostigma and 14 specimens from related groups (sections Adenophylli, Cupreati and Sampatik), recording and describing 45 qualitative leaf characters. These characters were assembled into a matrix and analysed using statistical clustering methods based on similarity. Our results show that trichomes are one of the most diverse and variable anatomical features among the studied specimens. Novel anatomical features for Croton include the presence of a hypodermis and two previously unreported types of non-glandular trichomes. Although we did not detect a single anatomical character uniting section Cyclostigma, combinations of anatomical characters were useful to establish species limits and taxonomic identities within this section.
Croton amentiformis, a new species of Euphorbiaceae from western South America, is described and illustrated. Morphological evidence and data from leaf anatomy indicate that the new species belongs to the dragon's blood group (Croton section Cyclostigma). The dense and congested inflorescence is unusual within the genus. This singular dragon's blood species is dedicated to the Florence Tropical Herbarium (FT) on the occasion of its 100th Anniversary celebrated in 2014.
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