The Boraginales are now universally accepted as monophyletic and firmly placed in Lamiidae. However, a consensus about familial classification has remained elusive, with some advocating recognition of a single, widely variable family, and others proposing recognition of several distinct families. A consensus classification is proposed here, based on recent molecular phylogenetic studies, morphological characters, and taking nomenclatural stability into consideration. We suggest the recognition of eleven, morphologically well-defined and clearly monophyletic families, namely the Boraginaceae s.str., Codonaceae, Coldeniaceae fam. nov., Cordiaceae, Ehretiaceae, Heliotropiaceae, Hoplestigmataceae, Hydrophyllaceae, Lennoaceae, Namaceae, and Wellstediaceae. Descriptions, synonomy, a taxonomic key, and a list of genera for these eleven families are provided, including the new family Coldeniaceae (monogeneric) and Namaceae (segregated from Hydrophyllaceae and comprising Nama, Eriodictyon, Turricula, and Wigandia), the latter necessitating a revised circumscription of a more morphologically coherent Hydrophyllaceae. Keywords angiosperms; Boraginaceae; Boraginales; classification; family; plant taxonomy
Boraginales Working Group • Families of Boraginales
503Version of Record TAXON 65 (3) • June 2016: 502-522 Boraginaceae in this traditional sense (Candolle, 1845; Bentham & Hooker, 1876;Gürke, 1893;Engler, 1898;Pilger & Krause, 1915) were subdivided into five subfamilies, namely Boraginoideae, Cordioideae, Ehretioideae, Heliotropioideae and Wellstedioideae. In pre-molecular times most scientists accepted this circumscription of Boraginaceae (e.g., Chadefaud & Emberger, 1960;Melchior, 1964b;Takhtajan, 1980Takhtajan, , 1997 Cronquist, 1981 Cronquist, , 1988Thorne, 1992), although some authors recognized one or the other subfamily at the family level. For example, Svensson (1925) andDi Fulvio (1978) removed Cordioideae, Heliotropioideae and Ehretioideae to Heliotropi aceae based on embryological studies, while Merxmüller (1960), Dahlgren (1980), and Takhtajan (1987) treated Wellstedioideae at the family level as Wellstediaceae. Conversely, Hoplestigmataceae, Hydrophyllaceae, and Lennoaceae were generally accepted as distinct families. However, the close relationships of these taxa to traditional Boraginaceae has been widely acknowledged by several authors (e.g., Jussieu, 1789; Baillon, 1891;Peter, 1893;Svensson, 1925; Chadefaud & Emberger, 1960; Melchior, 1964a, c;Takhtajan, 1980; Cronquist, 1981 Cronquist, , 1988. For example, Baillon (1891) defined the Boraginaceae as comprising nine series, which included both Boraginaceae and Hydrophyllaceae in their traditional circumscriptions. Chadefaud & Emberger (1960) considered Boraginaceae, Hoplestigmataceae, Hydrophyllaceae, and Lennoaceae to form a natural group within the order Tubiflorales. Takhtajan (1980) included these same families in the suborder Boraginineae.On the other hand, three groups historically associated to Boraginaceae have been clearly sho...