Hymenodictyeae (Rubiaceae – Cinchonoideae) and its phylogenetic significance

The tribe Naucleeae has recently been recircumscribed on the basis of both morphological and molecular [rbcL, trnT-F, internal transcribed spacer (ITS)] evidence, and has been found to be the sister group of the tribe Hymenodictyeae Razafim. & B. Bremer. In order to find pollen morphological support for this new classification, the pollen and orbicules of 65 species, representing 23 Naucleeae and the two Hymenodictyeae genera, were investigated by scanning electron and light microscopy. Naucleeae pollen is very small (< 20 µm) to small (20–30 µm) and its shape in equatorial view is suboblate to spheroidal or, more rarely, subprolate. Three compound apertures are present, each comprising a long and narrow ectocolpus, a circular to slightly lolongate mesoporus, and an often H-shaped endoaperture. The sexine ornamentation is perforate, rugulate, or (micro)reticulate, and supratectal elements are always absent. Apart from the variation in sexine ornamentation, the tribe is rather stenopalynous. The pollen of Hymenodictyeae is very similar to that of Naucleeae. The H-shaped endoapertures often observed probably form a synapomorphy for the clade comprising Naucleeae and Hymenodictyeae. Our pollen morphological observations are not in conflict with the widened delimitation of Naucleeae. Unambiguous pollen support for the recent subtribal or generic concepts of Naucleeae could not be found because of a lack of variation of pollen characters within the tribe. Orbicules are invariably present in the ten Naucleeae taxa investigated. They are spheroidal and smooth or irregularly folded.