Summary1. Functional traits are increasingly used to investigate community structure, ecosystem functioning or to classify species into functional groups. These functional traits are expected to be variable between and within species. Intraspecific functional variability is supposed to influence and modulate species responses to environmental changes and their effects on their environment. However, this hypothesis remains poorly tested and species are mostly described by mean trait values without any consideration of variability in individual trait values. 2. In this study, we quantify the extent of intraspecific plant functional trait variability, its spatial structure and its response to environmental factors. Using a sampling design structured along two direct and orthogonal climatic gradients in an alpine valley, we quantified and analysed the intraspecific variability for three functional traits (height, leaf dry matter content and leaf nitrogen content) measured on sixteen plant species with contrasting life histories. 3. Results showed a large variability of traits within species with large discrepancies between functional traits and species. This variability did not appear to be structured within populations. Between populations, the overall variability was partly explained by the selected gradients. Despite the strong effects of temperature and radiation on trait intraspecific variability, the response curves of traits along gradients were partly idiosyncratic. 4. Synthesis. Giving a comprehensive quantification of intraspecific functional variability through the analysis of an original data set, we report new evidence that using a single trait value to describe a given species can hide large functional variation for this species along environmental gradients. These findings suggest that intraspecific functional variability should be a concern for ecologists and its recognition opens new opportunities to better understand and predict ecological patterns in a changing environment. Further analyses are, however, required to compare inter-and intraspecific variability.

Summary1. The prevalence of phylogenetic niche conservatism (PNC) in nature is still a conflicting issue. Disagreement arises from confusion over its precise definition and the variety of approaches to measure its prevalence. Recent work highlighted that common measures of PNC strongly depend on the assumptions of the underlying model of niche evolution. However, this warning has not been well recognized in the applied literature and questionable approaches are still frequently applied. 2. The aim of this study is to draw attention to the assumptions underlying commonly applied simple measures of PNC. We used a series of simulations to illustrate how misleading results can be if assumptions of niche evolution are violated, that the violation of assumptions is a common phenomenon and that testing assumptions requires in-depth pretest. 3. We conclude that the seemingly simple measures of PNC, such as phylogenetic signal (PS) and evolutionary rate, are not so easy to apply if one accounts for the necessity to test model assumptions. In addition, these measures can be difficult to interpret. The common assumption that strong PS indicates PNC will be often invalid. In addition, the interpretation of some measures, for example the conclusion that evolutionary rate is slow enough to indicate PNC, requires a comparison with another clade, another trait or well-developed null model assumptions and thus additional data. 4. We suggest that studies investigating PNC should always compare alternative evolutionary models and that model comparisons should in particular include flexible niche evolution models such as multiple-optima OU models, although these are computational intensive. These models are directly inherited from the concept of macroevolutionary adaptive landscape and can indicate PNC either by relative few peak shifts or by narrow peaks in the adaptive landscape. A test of PNC thus requires comparing these parameters of the macroevolutionary landscape between clades or time periods. 5. The general prevalence of PNC in nature should be evaluated only based on studies keeping up to the high standards of communicating the used definition of PNC, testing the assumptions made in the modelling approaches and including newly developed models in a model comparison approach.

Relatively, few species have been able to colonize extremely cold alpine environments. We investigate the role played by the cushion life form in the evolution of climatic niches in the plant genus Androsace s.l., which spreads across the mountain ranges of the Northern Hemisphere. Using robust methods that account for phylogenetic uncertainty, intraspecific variability of climatic requirements and different life-history evolution scenarios, we show that climatic niches of Androsace s.l. exhibit low phylogenetic signal and that they evolved relatively recently and punctually. Models of niche evolution fitted onto phylogenies show that the cushion life form has been a key innovation providing the opportunity to occupy extremely cold environments, thus contributing to rapid climatic niche diversification in the genus Androsace s.l. We then propose a plausible scenario for the adaptation of plants to alpine habitats.

Aim Despite the accumulation of cases describing fast radiations of alpine plants, we still have limited understanding of the drivers of speciation in alpine floras and of the precise the timing of their diversification. Here, we investigated spatial and temporal patterns of speciation in three groups of alpine Primulaceae.Location Mountains of the European Alpine System. MethodsWe built a new phylogeny of Primulaceae including all species in three focal groups: Androsace sect. Aretia, Primula sect. Auricula and Soldanella. Combining phylogenetic information with a detailed climatic data set, we investigated patterns of range and ecological overlap between sister-species using an approach that takes phylogenetic uncertainty into account. Finally, we investigated temporal trajectories of diversification in the three focal groups.Results We found that a large majority of sister-species pairs in the three groups are strictly allopatric and show little differences in substrate and climatic preferences, a result that was robust to phylogenetic uncertainty. While rates of diversification have remained constant in Soldanella, both Androsace sect. Aretia and Primula sect. Auricula showed decreased diversification rates in the Pleistocene compared to previous geological epochs.Main conclusions Allopatric speciation with little niche divergence appears to have been by far the most common mode of speciation across the three groups studied. A few examples, however, suggest that ecological and polyploid speciation might also have played a role in the diversification of these three groups. Finally, extensive diversification likely occurred in the late Miocene and Pliocene coinciding with the later phases of the Alpine uplift, while diversification slowed down during subsequent glacial cycles of the Pleistocene.

Recent debate on whether climatic niches are conserved through time has focused on how phylogenetic niche conservatism can be measured by deviations from a Brownian motion model of evolutionary change. However, there has been no evaluation of this methodological approach. In particular, the fact that climatic niches are usually obtained from distribution data and are thus heavily influenced by biogeographic factors has largely been overlooked. Our main objective here was to test whether patterns of climatic niche evolution that are frequently observed might arise from neutral dynamics rather than adaptive scenarios. We develop a model inspired by Neutral Biodiversity Theory, where individuals disperse, compete, and undergo speciation independently of climate. We then sample the climatic niches of species according to their geographic position and show that even when species evolved independently of climate, their niches can nonetheless exhibit evolutionary patterns strongly differing from Brownian motion. Indeed, climatic niche evolution is better captured by a model of punctuated evolution with constraints due to landscape boundaries, two features that are traditionally interpreted as evidence for selective processes acting on the niche. We therefore suggest that deviation from Brownian motion alone should not be used as evidence for phylogenetic niche conservatism, but that information on phenotypic traits directly linked to physiology is required to demonstrate that climatic niches have been conserved through time.

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Aim-We still have limited understanding of the contingent and deterministic factors that have fostered the evolutionary success of some species lineages over others. We investigated how the interplay of intercontinental migration and key innovations promoted diversification of the genus Androsace.Location-Mountain ranges and cold steppes of the Northern Hemisphere.Methods-We reconstructed ancestral biogeographical ranges at regional and continental scales by means of a dispersal-extinction-cladogenesis analysis using dated Bayesian phylogenies and contrasting two migration scenarios. Based on diversification analyses under two frameworks, we tested the influence of life form on speciation rates and whether diversification has been diversitydependent.Results-We found that three radiations occurred in this genus, at different periods and on different continents, and that life form played a critical role in the history of Androsace. Shortlived ancestors first facilitated the expansion of the genus' range from Asia to Europe, while cushions, which appeared independently in Asia and Europe, enhanced species diversification in alpine regions. One long-distance dispersal event from Europe to North America led to the diversification of the nested genus Douglasia. We found support for a model in which speciation of the North American-European clade is diversity-dependent and close to its carrying capacity, and that the diversification dynamics of the North American subclade are uncoupled from this and follow a pure birth process. Europe PMC Funders Group Europe PMC Funders Author ManuscriptsEurope PMC Funders Author Manuscripts form was a convergent key innovation that fostered radiation into alpine habitats. Given the large ecological similarity of Androsace species, allopatry may have been the main mode of speciation.

The evolution of quantitative characters over long timescales is often studied using stochastic diffusion models. The current toolbox available to students of macroevolution is however limited to two main models: Brownian motion and the Ornstein-Uhlenbeck process, plus some of their extensions. Here, we present a very general model for inferring the dynamics of quantitative characters evolving under both random diffusion and deterministic forces of any possible shape and strength, which can accommodate interesting evolutionary scenarios like directional trends, disruptive selection, or macroevolutionary landscapes with multiple peaks. This model is based on a general partial differential equation widely used in statistical mechanics: the Fokker-Planck equation, also known in population genetics as the Kolmogorov forward equation. We thus call the model FPK, for Fokker-Planck-Kolmogorov. We first explain how this model can be used to describe macroevolutionary landscapes over which quantitative traits evolve and, more importantly, we detail how it can be fitted to empirical data. Using simulations, we show that the model has good behavior both in terms of discrimination from alternative models and in terms of parameter inference. We provide R code to fit the model to empirical data using either maximum-likelihood or Bayesian estimation, and illustrate the use of this code with two empirical examples of body mass evolution in mammals. FPK should greatly expand the set of macroevolutionary scenarios that can be studied since it opens the way to estimating macroevolutionary landscapes of any conceivable shape. [Adaptation; bounds; diffusion; FPK model; macroevolution; maximum-likelihood estimation; MCMC methods; phylogenetic comparative data; selection.].