Styloid crystals in Claoxylon (Euphorbiaceae) and allies (Claoxylinae) with notes on leaf anatomy Botanical Journal of the Linnean Society

Claoxylon and Micrococca are the only Euphorbiaceae genera that have rough dried leaves (fresh ones are smooth) because of protruding styloid (needle-like) crystals more or less perpendicular to the leaf surface, which perforate the epidermis and cuticle. A broad leaf anatomical study of the subtribes Claoxylinae, the monogeneric Lobaniliinae, and Mercurialis of the Mercurialinae (95 of a possible 235 species in all six genera) showed that styloids are present in Claoxylon, Discoclaoxylon, Erythrococca, Lobanilia, and Micrococca, and lacking in Claoxylopsis and Mercurialis. Contrary to Claoxylon, the dried leaves of Discoclaoxylon, Erythrococca, Lobanilia, and Micrococca are not rough, because the styloids do not perforate the epidermis during drying and therefore herbarium leaves generally remain smooth. The presence of styloids supports a clade in a recent molecular phylogenetic study that unites subtribes Claoxylinae and Lobaniliinae as Claoxylinae s.l. Mercurialis (no styloids) is sister to all other taxa (with styloids) in the monophyletic Claoxylinae s.l. The styloids form a synapomorphy for a monophyletic part of the Claoxylinae (Claoxylon, Discoclaoxylon, Erythrococca, Lobanilia, and Micrococca). Other leaf anatomical notes are provided, together with an overview of the occurrence of styloids, stomata, and druse crystals for most species in the Claoxylinae. The indumentum ranges from (virtually) absent to a sparse or dense tomentum of straight and thick- or thin-walled unicellular hairs (most common), curly hairs, or two-armed hairs (‘Malpighiaceous hairs’). The last two hair types probably form diagnostic characters for species groups within the genus Claoxylo