Sand-binding roots in Haemodoraceae: global survey and morphology in a phylogenetic context

To illustrate the morphology of sand-binding roots of Haemodoraceae, to conduct a comprehensive survey of the trait, spanning different climates across four continents, and to explore evolutionary hypotheses within a molecular phylogenetic framework. Sand-binding roots in Haemodoraceae were examined, measured and photographed in the field and on herbarium specimens. Photomicrographs were taken of southwest Australian species. The presence and absence of the sand-binding trait was mapped onto previously published phylogenies and an ancestral state reconstruction was performed. Sand grains were very tightly bound to the root surface by persistent root hairs in Haemodoraceae. The majority of genera and species were found to possess sand-binding roots and only 2 of the 14 genera, Conostylis and Tribonanthes, had sister taxa with and without the trait. The trait was recorded in tropical, sub-tropical and wet temperate species, but mainly in semi-arid species. Sand-binding roots were likely to have been present in the ancestor of the family and both sub-families. The presence of sand-binding roots is the probable ancestral condition for Haemodoraceae, associated with a high degree of phylogenetic conservatism and some secondary loss, notably in Conostylis. Experimental studies are needed to understand the ecological and evolutionary forces at work.