Species delimitation in Xanthium sect. Acanthoxanthium (Heliantheae, Asteraceae) and the neglected species Xanthium argenteum

Abstract

Xanthium is a genus of annual herbaceous plants that stands out within Asteraceae for being wind-pollinated, diclinous monoecious, and bearing solitary pistillate flowers in peculiar spiny structures (burs). Xanthium sect. Acanthoxanthium is native to South America and characterized by the presence of trifurcate spines at the base of the leaves. Past taxonomic treatments of the section have been contradictory, some recognising up to six species, others reducing all to a single polymorphic species. Altogether, 42 samples of X. sect. Acanthoxanthium were analysed, the vast majority taken from herbarium specimens between 20 and 160 years old. We sampled multiple specimens of X. spinosum from throughout its broad range as well as those taxa with narrower distributions, covering the whole range of morphological variation in the section. When possible, we included types and original material. We used Hyb-Seq techniques to obtain information from about 1000 single-copy nuclear genes and complete plastomes. Phylogenomic data were submitted to coalescent-based species delimitation approaches (SPEEDEMON). Additionally, we performed geometric morphometric analysis of leaf outlines. The results strongly support the identification of four lineages in the section favouring the acceptance of four of the hitherto described species, i.e., X. ambrosioides, X. argenteum, X. catharticum, and X. spinosum. These results were to some extent corroborated by morphometric analyses. While X. ambrosioides was well distinct from X. spinosum based on leaf morphology, such difference was not observed between X. spinosum and X. catharticum. However, X. catharticum differs from X. spinosum in its ecological requirements, being a species rather adapted to high-mountain environments of the Neotropics. Intriguingly, X. argenteum – a taxon described from a single herbarium collection – was also inferred as a species.