Distinctive wood anatomy of early-diverging Asteraceae: Barnadesioideae

Abstract

Abstract Asteraceae subfamily Barnadesioideae (ten genera, c. 90 species), confined to South America, are sister to the remainder of the family. The relative antiquity of the barnadesioids might lead one to expect that they contain more wood features plesiomorphic for the family, but only one character clearly falls in that category. Pits on imperforate tracheary elements are bordered (except for annuals), whereas simple pits occur in two related families, Calyceraceae (part) and Stylidiaceae (all that have been examined); in Goodeniaceae bordered pits only occur. By attaining fully bordered pits in Chuquiraga, the imperforate tracheary elements qualify as an apomorphy, ‘neotracheids’, valuable for resisting embolism formation in dry and cold South American habitats. Neotracheids are found also in Loricaria (Asteraceae: Inuleae), also from these habitats. Neotracheids, like plesiomorphic tracheids, are conductive, unlike fibre tracheids and libriform fibres. Other barnadesioid wood characters adapted to cold and drought include grouping of vessels, high vessel density, shorter vessel elements and helical sculpture (including helical thickenings on lumen-facing walls) of secondary xylem vessels. In Chuquiraga and Dasyphyllum, these helical thickenings are bordered in some species (new report for angiosperms). Some of the barnadesioid adaptations to cold and drought can be found in North American Artemisia spp. (Asteraceae: Anthemideae), especially in montane and desert areas. Wood features of barnadesioids match their respective habits and habitats: a few trees; shrubs of humid, dry or desert areas; a distinctive rhizomatous succulent in the pampas (Schlechtendalia); a scree/gravel perennial (Huarpea) and two genera of annuals, one with succulent leaves (Duseniella) and one with rayless (at least at first) stems in arid and open soils (Doniophyton). Diversity is unusual considering the small size of the subfamily. Examples of endodermal crystals (Arnaldoa only), pith sclereids and primary xylem fibres are cited.