The subfamily Mutisioideae (74 genera, ca. 865 species), which comprises three tribes, Mutisieae (43 genera, ca. 500 species), Nassauvieae (25 genera, ca. 315 species), and Stifftieae (six genera, 48 species), is analyzed at the generic level. A total of 87 genera traditionally related to Mutisioideae were studied. The genera of Mutisioideae are described and illustrated, and keys to tribes and genera are given. Corollas, anthers, styles, and pollen, are analyzed and discussed. The styles (smooth, rugulose to papillose, papillae short and rounded) constitute the main characters for delimiting the subfamily. The presence and distribution of the style papillae support the recognition of three tribes, although other characters as corolla shape contribute to their definition. The pollen of Nassauvieae can be clearly differentiated from the pollen of Mutisieae and Stifftieae. The morphology supports the exclusion of Brachylaena, and Warionia but not the exclusion of other taxa, for example, Hecastocleis, the Gochnatia complex, or the Ainsliaea group members.Resumen La subfamilia Mutisioideae (74 géneros, ca. 865 especies), que comprende tres tribus, Mutisieae (43 géneros, ca. 500 especies), Nassauvieae (25 géneros, ca. 315 especies), y Stifftieae (6 géneros, 48 especies), es analizada a nivel genérico. Se estudió un total de 87 géneros que tradicionalmente fueron relacionados a Mutisioideae. Se describen e ilustran los géneros de Mutisioideae, y se proveen claves para las tribus y los géneros. Se analizan y discuten las corolas, anteras y estilos. Los caracteres del estilo (glabro, ruguloso a papiloso, las papilas cortas y redondeadas) son los más importantes para delimitar la subfamilia. La presencia y distribución de las papilas del estilo ayudan al reconocimiento de las tres tribus, aunque otros caracteres como la forma de la corola contribuyen a su definición. El polen de Nassauvieae se puede diferenciar claramente del polen de Mutisieae y Stifftieae. La morfología apoya la exclusión de Brachylaena,

ABSTRACT. Onagraceae are a family of 17 genera in seven tribes, with the majority of species in tribes Onagreae and Epilobieae. Despite the species-richness of these two tribes, to date no phylogenetic study has been done with suf cient taxon sampling to examine relationships between and within these tribes. In this study, we used DNA sequence data from one nuclear region (ITS) and two chloroplast regions (trnL-trnF and rps16) to infer phylogenetic relationships among 93 taxa across the family, with concentrated sampling in the large tribe Onagreae. Results strongly suggest that tribe Gongylocarpeae is sister to tribes Epilobieae 1 Onagreae, both of which are monophyletic. Within Onagreae, Camissonia seems to be broadly paraphyletic, and Oenothera is also paraphyletic. In Oenothera there appear to be two lineages, one of which has Gaura 1 Stenosiphon nested within it. At the base of the Onagraceae phylogeny, we have clari ed previous confusion regarding con icting placements of Hauya and Lopezia based on nuclear versus chloroplast data. Results of these analyses are supported by morphology and suggest the need for new taxonomic delimitations, which are forthcoming.

The evolution of the inflorescence head in Asteraceae is important in the diversification of this largest angiosperm family. The aggregation of heads into higher-order capitulescences (secondary heads or syncephalia) is considered evolutionarily advanced. The genera Moscharia, Nassauvia, Polyachyrus, and Triptilion of the subtribe Nassauviinae (Mutisieae) have syncephalia with differing degrees of capitula condensation. ITS and plastid trnL-trnF regions were analyzed separately and together using maximum parsimony and maximum likelihood to examine the evolution of syncephalia in the Nassauviinae. The four genera displaying syncephalia do not form a clade minus taxa without syncephalia, indicating that secondary heads in Nassauviinae have either convergently evolved twice in the subtribe (or, very unlikely) once with multiple reversions. Strong support was obtained for a sister relationship between Leucheria (without syncephalium) and Polyachyrus, and both sister to Moscharia. Nassauvia and Triptilion form a distinct clade but are sister to other genera, Perezia and Panphalea, without syncephalium. Previous hypotheses postulated the evolution from simple to more complex secondary heads. We show that the ancestor of Moscharia, Polyachyrus, and Leucheria, in a more arid habitat, had a complex type of secondary head, and loss of complexity occurred in response to a shift from arid to mesic conditions.

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Fossil capitula and pollen grains of Asteraceae from the Eocene of Patagonia, southern Argentina, exhibit morphological features recognized today in taxa, such as Mutisioideae and Carduoideae, that are phylogenetically close to the root of the asteracean tree. This fossil supports the hypothesis of a South American origin of Asteraceae and an Eocene age of divergence and suggests that an ancestral stock of Asteraceae may have formed part of a geoflora developed in southern Gondwana before the establishment of effective dispersal barriers within this landmass.