Casuarinaceae are a Gondwanic family with a unique combination of morphological characters not comparable to any other family. Until recently, the 96 species in the family were classified in a single genus, Casuarina s.l. A recent morphological revision of the family resulted in the splitting of Casuarina s.l. into four genera-Allocasuarina, Casuarina s.s., Ceuthostoma, and Gymnostoma. This study uses matK sequence data from 76 species of Casuarinaceae and eight outgroup taxa to examine the phylogenetic structure within the Casuarinaceae. The study demonstrates the monophyly of the four genera and examines the relationships within the family; it tests the validity of the infra-generic subdivision of Allocasuarina; it discovers geography-based infra-generic subdivisions within Gymnostoma and Casuarina; and, finally, provides a molecular framework on which to trace the evolution of xeromorphy in the Casuarinaceae.