Molecular systematics and the origins of gypsophily in Nama L. (Boraginaceae)
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Nama L. is a genus of approximately 50 species of herbs and subshrubs that occurs in habitats ranging from arid deserts to mesic woodlands in the New World and the Hawaiian Islands. The group has historically been divided into five or six subgeneric groups based on habitat as well as on the morphology of the anthers, styles, leaves and seeds. At least 14 species of Nama from the Chihuahuan Desert Region are either facultatively or obligately endemic to gypsum deposits. This dissertation examines interspecies relationships within Nama from a molecular phylogenetic perspective in order to evaluate historic morphology-based subgeneric classification systems of the genus and to examine the origins of facultative and obligate gypsophily within the genus. DNA sequence data from the chloroplast regions matK and ndhF and from the nuclear ribosomal region ITS were collected from 46 species of Nama as well as from four new species and several outgroups. Data were analyzed using maximum likelihood and Bayesian methods. Phylogenetic analyses recover seven strongly supported major lineages within Nama. These lineages do not correspond to traditionally recognized subgenera, although they are largely congruent with an informal system based on ultrastructural observations of seeds. Four of the seven major lineages include gypsophilous species; these range from two lineages that include a single facultative gypsophile each, to one lineage that is almost entirely comprised of gypsophiles. Gypsum endemism in general, as well as facultative and obligate gypsophily in particular, has arisen multiple times in Nama. Parametric bootstrapping rejected the hypothetical monophyly of gypsophiles across the genus as a whole and within each of the two clades that contain multiple gypsophiles. Because approximately 20 species have been described since the last major revision of Nama nearly 80 years ago, detailed morphological observations of herbarium specimens were made in order to produce a comprehensive key to the species of Nama as well as the revision of a lineage comprising eight gypsophiles and one limestone endemic.