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Summary The evolution of secondary (insular) woodiness and the rapid disparification of plant growth forms associated with island radiations show intriguing parallels between oceanic islands and tropical alpine sky islands. However, the evolutionary significance of these phenomena remains poorly understood and the focus of debate. We explore the evolutionary dynamics of species diversification and trait disparification across evolutionary radiations in contrasting island systems compared with their nonisland relatives. We estimate rates of species diversification, growth form evolution and phenotypic space saturation for the classical oceanic island plant radiations – the Hawaiian silverswords and Macaronesian Echium – and the well‐studied sky island radiations of Lupinus and Hypericum in the Andes. We show that secondary woodiness is associated with dispersal to islands and with accelerated rates of species diversification, accelerated disparification of plant growth forms and occupancy of greater phenotypic trait space for island clades than their nonisland relatives, on both oceanic and sky islands. We conclude that secondary woodiness is a prerequisite that could act as a key innovation, manifest as the potential to occupy greater trait space, for plant radiations on island systems in general, further emphasizing the importance of combinations of clade‐specific traits and ecological opportunities in driving adaptive radiations.

The páramos, high-elevation Andean grasslands ranging from ca. 2800 m to the snow line, harbor one of the fastest evolving biomes worldwide since their appearance in the northern Andes 3–5 million years (Ma) ago. Hypericum (St. John's wort), with over 65% of its Neotropical species, has a center of diversity in these high Mountain ecosystems. Using nuclear rDNA internal transcribed spacer (ITS) sequences of a broad sample of New World Hypericum species we investigate phylogenetic patterns, estimate divergence times, and provide the first insights into diversification rates within the genus in the Neotropics. Two lineages appear to have independently dispersed into South America around 3.5 Ma ago, one of which has radiated in the páramos (Brathys). We find strong support for the polyphyly of section Trigynobrathys, several species of which group within Brathys, while others are found in temperate lowland South America (Trigynobrathys s.str.). All páramo species of Hypericum group in one clade. Within these páramo Hypericum species enormous phenotypic evolution has taken place (life forms from arborescent to prostrate shrubs) evidently in a short time frame. We hypothesize multiple mechanisms to be responsible for the low differentiation in the ITS region contrary to the high morphological diversity found in Hypericum in the páramos. Amongst these may be ongoing hybridization and incomplete lineage sorting, as well as the putative adaptive radiation, which can explain the contrast between phenotypic diversity and the close phylogenetic relationships.

BackgroundOur aim is to understand the evolution of species-rich plant groups that shifted from tropical into cold/temperate biomes. It is well known that climate affects evolutionary processes, such as how fast species diversify, species range shifts, and species distributions. Many plant lineages may have gone extinct in the Northern Hemisphere due to Late Eocene climate cooling, while some tropical lineages may have adapted to temperate conditions and radiated; the hyper-diverse and geographically widespread genus Hypericum is one of these.ResultsTo investigate the effect of macroecological niche shifts on evolutionary success we combine historical biogeography with analyses of diversification dynamics and climatic niche shifts in a phylogenetic framework. Hypericum evolved cold tolerance c. 30 million years ago, and successfully colonized all ice-free continents, where today ~500 species exist. The other members of Hypericaceae stayed in their tropical habitats and evolved into ~120 species. We identified a 15–20 million year lag between the initial change in temperature preference in Hypericum and subsequent diversification rate shifts in the Miocene.ConclusionsContrary to the dramatic niche shift early in the evolution of Hypericum most extant species occur in temperate climates including high elevations in the tropics. These cold/temperate niches are a distinctive characteristic of Hypericum. We conclude that the initial release from an evolutionary constraint (from tropical to temperate climates) is an important novelty in Hypericum. However, the initial shift in the adaptive landscape into colder climates appears to be a precondition, and may not be directly related to increased diversification rates. Instead, subsequent events of mountain formation and further climate cooling may better explain distribution patterns and species-richness in Hypericum. These findings exemplify important macroevolutionary patterns of plant diversification during large-scale global climate change.Electronic supplementary materialThe online version of this article (doi:10.1186/s12862-015-0359-4) contains supplementary material, which is available to authorized users.

Understanding how and why rates of evolutionary diversification vary is a key issue in evolutionary biology, ecology, and biogeography. Evolutionary rates are the net result of interacting processes summarized under concepts such as adaptive radiation and evolutionary stasis. Here, we review the central concepts in the evolutionary diversification literature and synthesize these into a simple, general framework for studying rates of diversification and quantifying their underlying dynamics, which can be applied across clades and regions, and across spatial and temporal scales. Our framework describes the diversification rate (d) as a function of the abiotic environment (a), the biotic environment (b), and clade‐specific phenotypes or traits (c); thus, d ~ a,b,c. We refer to the four components (a–d) and their interactions collectively as the “Evolutionary Arena.” We outline analytical approaches to this framework and present a case study on conifers, for which we parameterize the general model. We also discuss three conceptual examples: the Lupinus radiation in the Andes in the context of emerging ecological opportunity and fluctuating connectivity due to climatic oscillations; oceanic island radiations in the context of island formation and erosion; and biotically driven radiations of the Mediterranean orchid genus Ophrys. The results of the conifer case study are consistent with the long‐standing scenario that low competition and high rates of niche evolution promote diversification. The conceptual examples illustrate how using the synthetic Evolutionary Arena framework helps to identify and structure future directions for research on evolutionary radiations. In this way, the Evolutionary Arena framework promotes a more general understanding of variation in evolutionary rates by making quantitative results comparable between case studies, thereby allowing new syntheses of evolutionary and ecological processes to emerge.

Hypericum is a worldwide‐distributed genus with almost 500 species, including the medically used, facultative apomictic species H. perforatum. It is one of the few large plant genera for which alpha taxonomy has been completed and most species have been described. To conduct a formal cladistic analysis of the genus, we coded 89 morphological characters for all described taxa and analyzed the data for the species using parsimony and Bayesian methods. The obtained trees indicate that Hypericum is monophyletic if the monotypic genus Santomasia is included, and that Lianthus is its sister. The arrangement of the remaining genera of Hypericaceae included in the analysis is in congruence with molecular phylogenies. Within Hypericum the cladistic analysis revealed a basal grade containing Mediterranean species and three big clades containing most of the diversity of the genus. In contrast to earlier assumptions, we found no indication for an African origin of Hypericum, but assume that the genus evolved in what today is the Mediterranean area. Our phylogenies indicate a shrubby habit to be the ancestral state within Hypericum from which species with tree‐like and herbaceous habit evolved, and that apomixis originated at least three times independently within the genus.

Aim Tropical mountains around the world harbour an extraordinarily rich pool of plant species and are hotspots of biodiversity. Climatically, they can be zoned into montane climates at mid‐altitudes and tropical alpine climates above the tree line. Around half of the tropical alpine species belong to plant lineages with a temperate ancestry, although these regions are often geographically distant. We test the hypothesis that these temperate lineages are pre‐adapted to the tropical alpine climate. Location New World, with a focus on tropical alpine Andes. Time period Miocene to present. Major taxa studied Flowering plants. Methods We build multidimensional environmental models representing the full space of New World climates. We quantify the environmental similarity between the tropical alpine ecosystem and those of potential source areas, while correcting for regional differences by kernel density smoothers. Based on spatial observations of the genus Hypericum (St John's Wort), we quantify niche overlap and test for niche conservatism following intercontinental dispersal using density‐weighted nonparametric tests. A dated species tree, biogeographical estimation, multi‐optima Ornstein–Uhlenbeck models and model selection approaches are used to test for niche shifts during establishment in the tropical alpine Andes. Results The tropical alpine ecosystem is isolated by its climate from adjacent regions and is climatically similar to temperate lowland biomes of both hemispheres. Niche conservatism is evident in the study group, except in the tropical alpine lineage that is characterized by niche expansion and shifts in temperature optima. Main conclusions Our results reject the pre‐adaptation hypothesis and instead suggest pronounced niche evolution during colonization of tropical alpine ecosystems. Establishment involved substantial niche shifts, mainly in temperature‐related variables, and resulted in a tremendous proliferation of species in the newly invaded tropical alpine ecosystem.

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