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Premise of the Study: Floras of continental habitat islands, like those of islands, originate mostly through colonization, which can be followed by in situ speciation. We here address the question of the relative importance of colonization and in situ diversification in the high-altitude areas of the eastern African high mountains, the tropical Afroalpine Region, using the most species-rich genus in the region, Senecio, as an example., Methods: We expanded earlier Senecioneae phylogenies by adding more tropical African species and analyzed our phylogenetic tree biogeographically., Key Results: Senecio contains at least five clades with tropical African species, all of them containing tropical afroalpine species. Between four to 14 independent colonization events into the tropical Afroalpine most likely from montane regions in southern Africa were found. Additionally, relationships of tropical afroalpine species to Palearctic and South American taxa were identified. Although some in situ diversification occurred in Senecio in the tropical Afroalpine, the resulting number of species per clade is never higher than seven., Conclusion: Like other genera, Senecio colonized the tropical Afroalpine several times independently. Comparison with Mt. Kinabalu, a small tropical alpine-like region in Southeast Asia, and alpine-like regions in the Andes implies that rates of in situ speciation might be linked to area size., (© 2016 Botanical Society of America.)

Premise of the Study: Substrate specialization is often considered an important factor in evolutionary diversification. A classic example of divergence related to different substrate types is the dichotomy between calcicole and calcifuge plants on calcareous and siliceous substrates as found in the European Alps. When closely related species with contrasting substrate preferences are found in the same area, it is generally hypothesized that they diverged where they now occur. However, it is possible that Alpine edaphic diversity instead allows the coexistence of related species whose edaphic differentiation took place deeper in the phylogeny, in some other part of the range of their clades., Methods: We used sequences of the nuclear internal and external transcribed spacer regions to examine the origin of substrate differentiation in Minuartia series Laricifoliae, which contains many edaphic endemics, including a pair of Alpine taxa with contrasting substrate preferences: Minuartia langii (calcicole) and M. laricifolia (calcifuge)., Key Results: MINUARTIA LANGII and M. laricifolia are each more closely related to Balkan species than they are to each other and reached the Alps independently. The clade to which they belong is ancestrally calcicole. Minuartia langii inherited the ancestral ecology, while M. laricifolia is part of a subclade with serpentine endemics and one substrate generalist., Conclusions: In the study group, taxa with contrasting substrate preferences did not diverge in the Alps. Instead, taxa whose substrate differentiation arose elsewhere, likely on the Balkan Peninsula, were preadapted to take advantage of Alpine substrate diversity.

Premise of the Study: Heterogeneity of edaphic conditions plays a large role in driving the diversification of many plant groups. In the Alps and other European high mountains, many closely related calcicole and calcifuge plant taxa exist. To better understand patterns and processes of edaphic differentiation, the phylogeny of the edaphically variable genus Adenostyles was studied. The genus contains three species, of which A. alpina has five subspecies. Each species and subspecies is largely confined to either calcareous or noncalcareous substrates. •, Methods: We analyzed the phylogeny of Adenostyles using DNA sequences of nrITS, nrETS, nuclear chalcone synthase, and three plastid markers (rpl32-trnL, psbA-trnH, and ndhF-rpl32) from 45 in-group and five out-group samples. The phylogeny was used to reconstruct ancestral edaphic associations and distribution areas. •, Key Results: Within Adenostyles alpina, the shifts of edaphic association from calcicole to calcifuge in subsp. briquetii (Corsica) and in a clade of subsp. macrocephala (southernmost Italy) plus subsp. pyrenaica (Pyrenees) coincide with dispersal events. •, Conclusions: We conclude that colonization of areas with novel edaphic conditions via dispersal can trigger shifts of edaphic association. Accordingly, edaphic niche shifts can result from chance events.

Premise of the Study: The correct assessment of homology is an important prerequisite for reconstructing phylogenetic relationships and character evolution. Old World Papaveroideae (Papaver, Meconopsis, Roemeria, Stylomecon) show substantial diversity in gynoecium and capsule morphology. In particular, capsules can have distinct styles (Meconopsis p.p., Stylomecon) or a sessile stigmatic disc (Papaver). Molecular phylogenetic analyses of Old World Papaveroideae had shown that neither taxa with styles nor those with stigmatic discs represent monophyletic lineages. We here investigate whether either styles or stigmatic discs have arisen repeatedly during the diversification of Old World Papaveroideae., Methods: We investigated gynoecium ontogeny in Papaver rhoeas, P. californicum, Meconopsis cambrica, and Stylomecon heterophylla by scanning electron microscopy for the first time. Our observations were interpreted on the background of a well-resolved molecular phylogeny of the taxa investigated., Key Results: Papaver rhoeas and P. californicum share the presence of a developmentally complex garland-like stage in gynoecium ontogeny. The styles of M. cambrica and S. heterophylla result from growth in a ring-like zone beneath the carpel tips. This zone is also present in Papaver. In M. cambrica, traces of a garland-like stage can be seen. Style formation and stigma formation begin more or less simultaneously in M. cambrica, but style formation clearly precedes stigma formation in S. heterophylla., Conclusions: The styles present in M. cambrica and S. heterophylla are considered to have arisen in parallel from ancestors with a stigmatic disc. We speculate that style formation may have been a means to reduce selfing.

One of the longstanding questions in phylogenetic systematics is how to address incongruence among phylogenies obtained from multiple markers and how to determine the causes. This study presents a detailed analysis of incongruent patterns between plastid and ITS/ETS phylogenies of Tribe Senecioneae (Asteraceae). This approach revealed widespread and strongly supported incongruence, which complicates conclusions about evolutionary relationships at all taxonomic levels. The patterns of incongruence that were resolved suggest that incomplete lineage sorting (ILS) and/or ancient hybridization are the most likely explanations. These phenomena are, however, extremely difficult to distinguish because they may result in similar phylogenetic patterns. We present a novel approach to evaluate whether ILS can be excluded as an explanation for incongruent patterns. This coalescence-based method uses molecular dating estimates of the duration of the putative ILS events to determine if invoking ILS as an explanation for incongruence would require unrealistically high effective population sizes. For four of the incongruent patterns identified within the Senecioneae, this approach indicates that ILS cannot be invoked to explain the observed incongruence. Alternatively, these patterns are more realistically explained by ancient hybridization events.

To explore the biogeographic history of Mediterranean/arid plant disjunctions, Old and New World Senecio sect. Senecio were analyzed phylogenetically using nuclear ribosomal DNA sequences (ITS). A clade corresponding to sect. Senecio was strongly supported. Area optimization indicated this clade to be of southern African origin. The Mediterranean and southern African floras were not distinguishable as sources of the main New World lineage, estimated to have become established during the middle Pliocene. Another previously suspected recent dispersal to the New World from the Mediterranean was confirmed for the recently recognized disjunction in S. mohavensis. The loss of suitable land connections by the Miocene means that both New World lineages must represent long-distance dispersal, providing the first evidence of repeat intercontinental dispersal in a Mediterranean group. In contrast, migration within Africa may have utilized an East African arid corridor. Recent dispersal to northern Africa is supported for S. flavus, which formed part of a distinct southern African lineage. Novel pappus modifications in both disjunct species may have enabled dispersal by birds. An estimated early Pliocene origin of sect. Senecio coincides with the appearance of summer-dry climate. However, diversification from 1.6 BP highlights the importance of Pleistocene climate fluctuations for speciation.

Soldanella contains 16 species of herbaceous perennials that are endemic to the central and south European high mountains. The genus is ecogeographically subdivided into forest/montane and alpine species. Evolutionary relationships and large-scale biogeographic patterns were inferred from parsimony analyses of the internal transcribed spacer (ITS) regions of nuclear ribosomal DNA, and genetic distance analyses based on amplified fragment length polymorphism (AFLP) markers. The ITS region proved useful for examining subgeneric relationships and testing hypotheses on genus-wide divergence times, whereas the AFLP markers were suitable for studying relationships among closely related taxa and biogeographic patterns of divergence. Neither ITS nor AFLP data supported sectional delimitations, particularly those related to the grouping of S. alpina (sect. Soldanella) with S. pusilla (sect. Tubiflores), which may be the result of hybridization. Additional results and conclusions drawn are (1) Soldanella is derived from an ancestor of Asian origin with a montane ecology; (2) estimates of divergence times suggest a late Quaternary origin of the genus; (3) alpine species of sect. Tubiflores diverged from within a paraphyletic sect. Soldanella of mainly montane species; (4) alpine and montane species of Soldanella experienced different cycles of range expansion and contraction during late Quaternary climatic changes, resulting in differential patterns of geographic distribution; and (5) AFLP divergence among montane species from eastern Europe was lower than between alpine species; we hypothesize that the latter differentiated in allopatric regions of expansion during glacials, while the former experienced secondary contact at lower elevations in more southern refugia.