No abstract

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,

Coihimtia isowv of llic largest ^t'lit'ia of \\\c Irilx' Mutisieae (Aslcraceac) ami has IxM'ri Iradilionally charactrri/rd by its honio^anious rupitula wilh isoinuqiliu-toiollas. A itKuplioloi^ical sluil} o[ GocluKtlid and associated ^t'lu'ia. i.e., Arfinosrris, Chnvoa, (Inicothdiunus, ('yrlolcpis, Uyalis, lauthopappiis, Nonelia, Plciotaxis, and Wundvrluhia, was canlcd out lo rvalna!<' \\\v circiiinscriplion (»f (Unhnutid and ils sections, and the affitiilirs of this complex of genera. The characters studied involve hahil, U^af features (consistency, |>ul)escence lyj>e, shape, margin, and venation), types of capiluiescence, inxolu

Subfamily Gochnatioideae is the sister group of ∼96% of the species in Compositae (Asteraceae). It is of particular interest not only because of its position in the phylogeny, but also because, in recent molecular studies, the node it occupies is not strongly supported making difficult any inferences on the direction of character evolution in the family. The recognition of tribe Gochnatieae was one of the results of a comprehensive molecular analysis of the family that showed the traditional circumscription of the Mutisieae to be non–monophyletic. The four genera of Gochnatieae (Cnicothamnus, Cyclolepis, Gochnatia, Richterago) were defined by the presence of apiculate anther appendages and dorsally smooth style branches. Gochnatia, which contained about 70 species, was the largest and most complex genus and in the last decade some of its sections have been moved (or returned) to the rank of genus. This study includes a large selection of potential outgroups and over 60% of all species in the tribe, including all the genera and all but one of the sections of Gochnatia, to examine evolutionary relationships among the taxa. Both cpDNA and nrDNA were used in a phylogenetic analysis using parsimony, likelihood, and Bayesian approaches. The results suggest a non–monophyletic Gochnatia that is here resolved by the recognition of segregate genera. Morphological characters support these new genera and allow the adoption of a new classification for the Gochnatieae. A biogeographic analysis shows a possible southern South American/Andean origin followed by movement in three directions: into the Central Andes, into central and northern Brazil, and into Mexico and the Caribbean. The dating analysis gives an age of the split of the core Gochnatieae from the Wunderlichieae–Cyclolepis clade, and hence the age of the tribe, of 36–45 Ma and an age of 23–25 Ma for the first split within the core Gochnatieae (Andean vs. Brazil–Mexico–Caribbean). Cnicothamnus remains in Gochnatieae but Cyclolepis is designated incertae sedis.

The new genus Moquiniastrum, the result of recent phylogenetic analyses, is described. Although these analyses are based on cpDNA and nDNA, they also involve documentation of the distinctive morphological characters supporting this new genus. The recognition of Moquiniastrum is necessary to accurately reflect the relationships of the taxa found in the tribe Gochnatieae. Moquiniastrum includes twenty-one species that are usually gynodioecious and found mainly in Brazil but with some species elsewhere in South America. A description of Moquiniastrum, together with the corresponding new combinations, new lectotypifications of three names and one new neotypification is here provided.

In the context of recent molecular phylogenies of the basal grades of Compositae, we investigated the utility of pollen morphology within the tribe Gochnatieae. The pollen of 64 species of Anastraphia, Cnicothamnus, Cyclolepis, Gochnatia, Pentaphorus, and Richterago was studied using light microscopy and scanning electron microscopy. In addition, three extra-Gochnatieae genera (Ianthopappus, Leucomeris, and Nouelia) were examined as they were traditionally morphologically related to members of the tribe Gochnatieae. Three of the species of Gochnatieae were examined using transmission electron microscopy. Two pollen types, and two new subtypes, have been recognized on the basis of the pollen shape, size, and exine sculpture. The pollen features of Gochnatia sect. Moquiniastrum and G. cordata are similar and distinctive within the genus and support the recently re-circumscribed section Hedraiophyllum. Within the species with echinate pollen surface, the distinctive spine length of Anastraphia supports its recent resurrection as a genus. The identity of Pentaphorus could not be supported by pollen features as was for other morphological characteristics. The pollen features shared across Cyclolepis, Ianthopappus, Leucomeris, Nouelia and Gochnatia sect. Moquiniastrum, as well as those shared by Richterago and Anastraphia could be a result of parallel evolution.

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Smallanthus (Asteraceae, Millerieae), the “yacón” genus, comprises 23 species ranging from southeastern United States to central Argentina, with the greatest diversity in Peru and Mexico. Species of Smallanthus are herbs, shrubs or small trees with tuberous roots, opposite, trinerved or triplinerved leaves, paleate receptacle, ray corollas externally hairy at the base, pappus lacking, and marginal cypselae partially embraced by the inner phyllaries of the involucre. As a result of this revision: Smallanthus araucariophilus and S. riograndesis are considered synonyms of S. macroscyphus; lectotypes are designated for nine names; and the geographical distribution of S. latisquamus, S. lundelli, S. riparius, and S. siegesbeckia is extended. A comprehensive key to the species and detailed descriptions, geographical distribution, phenology, illustrations and distribution maps are provided for all taxa.