Phytotaxa 159 (2): 077–094 www.mapress.com / phytotaxa / Copyright © 2014 Magnolia Press ISSN 1179-3155 (print edition) Article PHYTOTAXA ISSN 1179-3163 (online edition) http://dx.doi.org/10.11646/phytotaxa.159.2.2 Phylogenetic studies in Smallanthus (Millerieae, Asteraceae): a contribution from morphology MAIRA S. VITALI1 & JESSICA N. VIERA BARRETO1 1 División Plantas Vasculares, Museo de La Plata, Paseo del Bosque s/n, 1900 La Plata, Argentina E-mail: vitali@fcnym.unlp.edu.ar (corresponding author). E-mail: jvierabarreto@fcnym.unlp.edu.ar Abstract We present a cladistic analysis of all the species of Smallanthus. Six taxa within Rumfordia, Ichthyothere, Acanthospermum and Tridax served as outgroups. We evaluated the monophyly and the relationships between the species of Smallanthus through a maximum parsimony study based on morphological data. The matrix included 31 qualitative characters from floral and vegetative parts of the specimens. We also explored the phylogenetic significance of treating quantitative characters as continuous. Only one most parsimonious tree was obtained. In agreement with previous phylogenetic studies based on molecular data, we recovered a monophyletic Smallanthus. The presence of ray corollas, densely pubescent at the base, was the synapomorphy that defined Smallanthus. Smallanthus microcephalus and two other major clades were recovered. The first clade included S. glabratus, S. fruticosus, S. jelskii and S. pyramidalis, while the second one contained the remaining species of Smallanthus. The analysis recovered one species of Rumfordia as sister to Smallanthus. We present a new combination, Smallanthus cocuyensis, based on morphological analysis of the type specimen. Keywords: Cladistic analysis, Compositae, quantitative characters, Rumfordia Resumen En este trabajo se presenta un análisis cladístico de todas las especies del género Smallanthus. Se utilizaron como grupos externos seis especies potencialmente relacionadas incluidas en los géneros Rumfordia, Ichthyothere, Acanthospermum y Tridax. Se evaluó la monofilia y relaciones entre las especies de Smallanthus a través de un estudio de máxima parsimonia basado en caracteres morfológicos. La matriz incluyó treinta y un caracteres cualitativos, tanto florales como vegetativos. También se analizó la importancia filogenética de los caracteres cuantitativos tratados como continuos. Se obtuvo como resultado un único árbol más parsimonioso. Como resultado se obtuvo al género Smallanthus como monofilético, apoyando estudios previos basados en datos moleculares. La presencia de una densa concentración de pelos en la base de las corolas de las flores del radio fue la sinapomorfía que soportó a Smallanthus. Dentro de Smallanthus se obtuvieron dos grandes clados y un linaje independiente, Smallanthus microcephalus. El primer clado incluyó a S. glabratus, S. fruticosus, S. jelskii y S. pyramidalis, y el segundo se encontró formado por las restantes especies de Smallanthus. El análisis mostró a una especie de Rumfordia como hermana del género Smallanthus. Se propone una nueva combinación, Smallanthus cocuyensis, en base a observaciones basadas en la morfología del material tipo. Introduction Smallanthus Mackenzie (Asteraceae, Millerieae) includes 24 species distributed from the southern United States to Central-East Argentina. Many species of Smallanthus occur in Mexico (seven species) and Peru (eight species). The remaining species are distributed in North America (United States), Central America (Costa Rica, El Salvador, Honduras, Nicaragua and Panama) and South America (Argentina, Bolivia, Brazil, Colombia, Ecuador, Paraguay, Accepted by Alexander Sennikov: 10 Jan. 2014; published: 11 Feb. 2014 77 Conclusion Based on the results of our single most parsimonious tree we could consider Smallanthus as monophyletic, in agreement with previous molecular analyses. Smallanthus is defined here by ray corollas densely pubescent at the base. We could recognize two clades in Smallanthus, the first including S. glabratus, S. fruticosus, S. jelskii and S. pyramidalis, and the second including S. siegesbeckius, S. macroscyphus, S. maculatus, S. riparius, S. uvedalius, S. cocuyensis, S. meridensis, S. oaxacanus, S. mcvaughii, S. sonchifolius, S. parviceps, S. riograndensis, S. apus, S. latisquamus, S. quichensis, S. lundelli, S. obscurus, S. putlanus and S. araucariophilus. These clades, however, had a weak support. The species of the first clade share several morphological characters with Rumfordia. Furthermore, as a contribution to the understanding of the relationships between subtribes of Millerieae, Ichthyothere was recovered as phylogenetically more distantly related to Smallanthus than to Rumfordia. Based on our results, Polymnia cocuyensis and Smallanthus meridensis were regarded as distinct entities. Regarding the characters used for this study, we could conclude that: 1) quantitative characters were found to be informative when treated as continuous variables, 2) the presence of functionally staminate disc florets provided no information on the groups analyzed in our work, and neither did the bilabiate-like marginal corollas. The evidence from chemical studies on species of Smallanthus with medicinal uses partially support some of the clades obtained in our analyses. The major clades of Smallanthus contained both South American and Central or North American species, suggesting that dispersal across the Isthmus of Panama has occurred multiple times in the history of the genus, as stated by other studies on this tribe. Further studies at the level of subtribe, based on both molecular and morphological characters, would be necessary to resolve questions about the relationship of Smallanthus with other genera of the subtribe Millerieae, particularly with the potentially closely related genus Rumfordia. Acknowledgements We are grateful to Alexander Sennikov, the editor, and the reviewers who provided valuable comments on an earlier draft of this paper and to the curators of herbaria for lending specimens and sharing digital photographs. We thank Gisela Sancho for her invaluable help and for providing critical discussion throughout the production of this manuscript. 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