The Lactucinae or Lactuca alliance include approximately 200 species distributed across the Northern Hemisphere and Africa. They were not recognised as a separate lineage until the late 20th century and their circumscription is still not fully settled. The generic classification of no other group of the Cichorieae has faced as many controversies as the Lactuca alliance and competing taxonomies coexist. This paper provides the first molecular phylogeny of the subtribe on a global scale as a major step towards its revised systematics. The sampling includes almost 60% of the species-level diversity and spans all species groups. Two datasets were created, one including the nrDNA ITS region, the other five concatenated non-coding plastid DNA loci. Maximum parsimony, maximum likelihood and Bayesian inference were used to produce a robust phylogenetic backbone. The diversification and expansion of the Lactucinae in a geohistorical context was reconstructed by estimating the age of their lineages using relaxed molecular clock dating and by inferring the ancestral areas using Bayesian binary analysis. The redelimited monophyletic Lactucinae are composed of seven lineages that also include Prenanthes, which is confirmed to have a single species, P. purpurea. The positions of two further lineages shift between Lactucinae and Crepidinae in the nuclear and plastid DNA phylogenies. Incongruence between the phylogenies suggests events of ancient reticulation or incomplete lineage sorting in the formation of these latter two lineages and in two of the seven other Lactucinae lineages. The phylogenetic results show a dilemma for Lactucinae systematics: most generic concepts proposed to date are highly artificial but the resolved phylogenetic lineages do not constitute practicable taxonomic entities with our current knowledge. Diversification of the subtribe is inferred to have taken place since the Middle Miocene. Biogeographic analysis proposes that the clade originated in the mountains of the landmass mediating between the European and Asian continents and delimited in the south by the Tethys Sea and in the north by the Paratethys Sea. Several independent migrations have occurred into various parts of Asia, Europe, tropical Africa and North America.

The first comprehensive molecular phylogenetic reconstruction of the Cichorieae subtribe Lactucinae is provided. Sequences for two datasets, one of the nuclear rDNA ITS region, the other of five concatenated non-coding chloroplast DNA markers including the petD region and the psbA-trnH, 5′trnL(UAA)-trnF, rpl32-trnL(UAG) and trnQ(UUG)-5′rps16 spacers, were, with few exceptions, newly generated for 130 samples of 78 species. The sampling spans the entire subtribe Lactucinae while focusing on its Chinese centre of diversity; more than 3/4 of the Chinese Lactucinae species are represented. The nuclear and plastid phylogenies inferred from the two independent datasets show various hard topological incongruences. They concern the internal topology of major lineages, in one case the placement of taxa in major lineages, the relationships between major lineages and even the circumscription of the subtribe, indicating potential events of ancient as well as of more recent reticulation and chloroplast capture in the evolution of the subtribe. The core of the subtribe is clearly monophyletic, consisting of the six lineages, Cicerbita, Cicerbita II, Lactuca, Melanoseris, Notoseris and Paraprenanthes. The Faberia lineage and the monospecific Prenanthes purpurea lineage are part of a monophyletic subtribe Lactucinae only in the nuclear or plastid phylogeny, respectively. Morphological and karyological support for their placement is considered. In the light of the molecular phylogenetic reconstruction and of additional morphological data, the conflicting taxonomies of the Chinese Lactuca alliance are discussed and it is concluded that the major lineages revealed are best treated at generic rank. An improved species level taxonomy of the Chinese Lactucinae is outlined; new synonymies and some new combinations are provided.

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