MORPHOLOGY AND PHYLOGENETIC INTERRELATIONSHIPS OF THE ASTERACEAE, CALYCERACEAE, CAMPANULACEAE, GOODENIACEAE, AND RELATED FAMILIES (ASTERALES)

In search for the sister group of the Asteraceae, morphological evidence was assembled for investigating the relationship between the Asteraceae and those families most frequently considered to be their closest relatives, in particular the Calyceraceae, Campanulaceae (along with the frequently included Lobeliaceae, Cyphiaceae, Cyphocarpaceae, and Nemacladaceae), and Goodeniaceae (and the sometimes included Brunoniaceae). Several other families that have been associated with this group of families, the ''Asterales-Campanulales-complex,'' were also considered: Pentaphragmataceae, Sphenocleaceae, Stylidiaceae, Donatiaceae, Menyanthaceae, and Argophyllaceae. In order to delineate the complex more precisely, another eight putatively related families were also included in the analysis. Cladistic parsimony analysis of 46 morphological and chemical characters for the 23 families was undertaken. Stability of the branches was estimated by the number of extra steps necessary to lose the group, as well as by the number of reweighted extra steps (using rescaled consistency indices) necessary to lose the group (a new approach). The results indicate that there is a monophyletic group of 14 families comprising those of the Asterales-Campanulales-complex as well as Pentaphragmataceae, Sphenocleaceae, Stylidiaceae, Donatiaceae, and Menyanthaceae; this group is recognized as the order Asterales. Within the order, the Asteraceae, Calyceraceae, Brunoniaceae, and Goodeniaceae form one comparatively well-supported clade and the five families of the Campanulaceae sensu late form another well-supported clade.