Labile Trait Research Articles

The influence of pre‐exposure to marine heatwaves on the critical thermal maxima (CTmax) of marine foundation species

Abstract Marine foundation species underpin some of the world's most diverse ecosystems but they are increasingly threatened by intensification of marine heatwaves (MHWs). Where MHWs exceed critical thermal maxima (CTmax), increased mortality and population declines can occur. CTmax is increasingly used to assess MHW population vulnerability but studies estimating CTmax across species, range edges and thermal histories in a comparable manner remain lacking. We determined the impact of MHWs on subsequent CTmax estimates of matched cool/warm affinity pairs of marine foundation species (kelp, seagrass and bivalves) in the Western English Channel. Following a 4‐week MHW simulation, individuals were subjected to a CTmax trial, where temperatures were raised by 2°C day−1 until physiological end points were reached. We found no positive effect of MHWs on CTmax but clear negative impacts were observed for some groups of foundation species. Increased MHW intensity had a stepwise negative impact on the physiology of both warm (Laminaria ochroleuca) and cool water (L. digitata) kelp species that manifested in significant reductions in CTmax. Surprisingly, this was most marked in the warm water species, which runs opposite to the assumed safety of leading‐edge populations. The physiology of warm (Zostera noltii) and cool (Z. marina) seagrasses was negatively impacted by increasing MHW intensity but no significant decrease in CTmax was observed. Both bivalve species (Mytilus edulis and Magallana gigas) showed marked resistance to exposure to MHWs, which was unexpected given the observed vulnerability of these species to stressful summertime conditions. Our results show pre‐exposure to realistic MHWs can influence CTmax values but generalities are difficult to make across groups or based on assumed thermal safety margins. We show CTmax is a labile trait and exposure to MHWs, can erode the resilience of an individual or population to subsequent thermal challenges. This leaves uncertainty within frameworks built to understand where and when MHWs will be most impactful. Further experimentation across a wider range of species and thermal challenges is needed to better understand the dynamic nature of CTmax and field validation is needed to determine the responses of individuals and populations within complex natural systems. Read the free Plain Language Summary for this article on the Journal blog.

Consistent differences in tissue oxygen levels across 15 insect species reflect a balance between oxygen supply and demand and highlight a hitherto unknown adaptation for extracting sufficient oxygen from water

Animals, including insects, need oxygen for aerobic respiration and eventually asphyxiate without it. Aerobic respiration, however, produces reactive oxygen species (ROS), which contribute to dysfunction and aging. Animals appear to balance risks of asphyxiation and ROS by regulating internal oxygen relatively low and stable, but sufficient levels. How much do levels vary among species, and how does variation depend on environment and life history? We predicted that lower internal oxygen levels occur in insects with either limited access to environmental oxygen (i.e., insects dependent on aquatic respiration, where low internal levels facilitate diffusive oxygen uptake, and reduce asphyxiation risks) or consistently low metabolic rates (i.e., inactive insects, requiring limited internal oxygen stores). Alternatively, we predicted insects with long life-stage durations would have internal oxygen levels > 1 kPa (preventing high ROS levels that are believed to occur under tissue hypoxia). We tested these predictions by measuring partial pressures of oxygen (PO2) in tissues from juvenile and adult stages across 15 species comprising nine insect orders. Tissue PO2 varied greatly (from 0 to 18.8 kPa) and variation across species and life stages was significantly related to differences in habitat, activity level, and life stage duration. Individuals with aquatic respiration sustained remarkably low PO2 (mean = 0.88 kPa) across all species from Ephemeroptera (mayflies), Plecoptera (stoneflies), Trichoptera (caddisflies), and Diptera (true flies), possibly reflecting a widespread, but hitherto unknown, adaptation for extracting sufficient oxygen from water. For Odonata (dragonflies), aquatic juveniles had higher PO2 levels (mean = 6.12 kPa), but these were still lower compared to terrestrial adults (mean = 13.3 kPa). Follow-up tests in juvenile stoneflies showed that tissue PO2 remained low even when exposed to hyperoxia, suggesting that levels were down-regulated. This was further corroborated since levels could be modulated by ambient oxygen levels in dead individuals. In addition, tissue PO2 was positively related to activity levels of insect life stages across all species and was highest in stages with short durations. Combined, our results support the idea that internal PO2 is an evolutionarily labile trait that reflects the balance between oxygen supply and demand within the context of the environment and life-history of an insect.

Individual variation in thermally induced plasticity of metabolic rates: ecological and evolutionary implications for a warming world.

Energy metabolism is a fundamental property of life providing the energy for all processes and functions within an organism. As it is temperature-dependent, it mediates the effects of changing climate on ectotherm fitness and population dynamics. Though resting metabolic rate is a highly labile trait, part of its variation is individually consistent. Recent findings show that resting metabolic rate contains consistent variation not only in the elevations (intercepts) but also in the slopes of individual thermal dependence curves, challenging the thermal dependence assumption for this trait in several ectotherm taxa. I argue that among-individual variation in thermal metabolic curves represents a previously undetected component of ectotherm response to climate change, potentially affecting their adaptive capacity and population resilience under increasing stochasticity of thermal environment. Future studies need to examine not only the amount of among-individual variation in thermal metabolic curves across phylogenetic contexts but also other aspects concerning its mechanisms and adaptive significance to improve predictions about the impact of climate change on ectotherm population dynamics. This article is part of the theme issue 'The evolutionary significance of variation in metabolic rates'.

Ploidy evolution in a wild yeast is linked to an interaction between cell type and metabolism

Ploidy is an evolutionarily labile trait, and its variation across the tree of life has profound impacts on evolutionary trajectories and life histories. The immediate consequences and molecular causes of ploidy variation on organismal fitness are frequently less clear, although extreme mating type skews in some fungi hint at links between cell type and adaptive traits. Here, we report an unusual recurrent ploidy reduction in replicate populations of the budding yeast Saccharomyces eubayanus experimentally evolved for improvement of a key metabolic trait, the ability to use maltose as a carbon source. We find that haploids have a substantial, but conditional, fitness advantage in the absence of other genetic variation. Using engineered genotypes that decouple the effects of ploidy and cell type, we show that increased fitness is primarily due to the distinct transcriptional program deployed by haploid-like cell types, with a significant but smaller contribution from absolute ploidy. The link between cell-type specification and the carbon metabolism adaptation can be traced to the noncanonical regulation of a maltose transporter by a haploid-specific gene. This study provides novel mechanistic insight into the molecular basis of an environment–cell type fitness interaction and illustrates how selection on traits unexpectedly linked to ploidy states or cell types can drive karyotypic evolution in fungi.

The evolutionary ecology of variation in labile traits: selection on its among- and within-individual components.

Closer integration between behavioral ecology and quantitative genetics has resulted in a recent increase in studies partitioning sources of variation in labile traits. Repeatable between-individual differences are commonly documented, and their existence is generally explained using adaptive arguments, implying that selection has shaped variation at the among- and within-individual level. However, predicting the expected pattern of non-adaptive phenotypic variation around an optimal phenotypic value is difficult, hampering our ability to provide quantitative assessments of the adaptive nature of observed patterns of phenotypic variation within a population. We argue that estimating the strength of selection on trait variation among and within individuals provides a way to test adaptive theory concerned with phenotypic variation. To achieve this aim, we describe a nonlinear selection analysis that enables the study of the selective pressures on trait means and their among- and within-individual variation. By describing an integrative approach for studying the strength of selection on phenotypic variation at different levels, we hope to stimulate empirical studies investigating the ecological factors that can shape the repeatability, heritability, and coefficients of variation of labile and other repeatedly expressed traits.

Morphological trait evolution inSolanum(Solanaceae): Evolutionary lability of key taxonomic characters

AbstractSolanumis one of the world's largest and economically most important plant genera, including 1245 currently accepted species and several major and minor crops (e.g., tomato, potato, brinjal eggplant, scarlet eggplant, Gboma eggplant, lulo, and pepino). Here we provide an overview of the evolution of 25 key morphological traits for the major and minor clades of this giant genus based on stochastic mapping using a well‐sampled recently published phylogeny ofSolanum. The most evolutionarily labile traits (showing >100 transitions across the genus) relate to plant structure (growth form and sympodial unit structure), herbivore defence (glandular trichomes), pollination (corolla shape and colour), and dispersal (fruit colour). Ten further traits show evolutionary lability with 50–100 transitions across the genus (e.g., specialised underground organs, trichome structure, leaf type, inflorescence position and branching, stamen heteromorphism). Our results reveal a number of highly convergent traits inSolanum, including tubers, rhizomes, simple leaves, yellow corollas, heteromorphic anthers, dioecy, and dry fruits, and some unexpected pathways of trait evolution that could be explored in future studies. We show that informally named clades ofSolanumcan be morphologically defined by trait combinations providing a tool for identification and enabling predictive phylogenetic placement of unsampled species.

Contrasting patterns in phylogenetic and biogeographic factories of invasive grasses (Poaceae) across the globe

Grasses (Family Poaceae) are among the most successful invasive plants in the world. Here we evaluate phylogenetic and biogeographic patterns of emergence of naturalized and invasive species among grasses globally. In our data, circa 19% of the grasses are currently catalogued as invasive and almost 38% are listed as naturalized; these are among the highest ratios for single families of organisms. Remarkably, most tribes of grasses contain numerous naturalized and invasive species, suggesting that the invasion success is rooted broadly in ancestral traits in the Poaceae. Moreover, the probability of invasiveness is positively related to the diversification rates in the family also suggesting a link with recent radiation events. The phylogenetic distribution of the invasive condition is neither strongly conserved nor purely random. Phylogenetic clumping levels also vary between Poaceae subclades. We postulate that this diffuse clumping could be partially attributed to the expression of labile traits that contribute to species invasiveness. In addition, floristic regions (biomes and biogeographic realms) have different proportions of invasive species, with the temperate Palearctic region having the highest ratio of invasive vs. non-invasive species. The phylodiversity of aliens across regions is also variable in space. Comparison of alien phylodiversity levels across biogeographic realms and biomes reveals regions producing highly restricted invasive lineages and others where the diversity of aliens exported is no different from global mean diversity levels in grasses. Elucidating the evolutionary patterns and drivers of invasiveness is useful for understanding and managing invasions, with the low phylogenetic structure of alien grasses warning of their overall high invasiveness potential.

The evolution of early-life telomere length, pace-of-life and telomere-chromosome length dynamics in birds.

Telomeres, the short DNA sequences that protect chromosome ends, are an ancient molecular structure, which is highly conserved across most eukaryotes. Species differ in their telomere lengths, but the causes of this variation are not well understood. Here, we demonstrate that mean early-life telomere length is an evolutionary labile trait across 57 bird species (representing 35 families in 12 orders) with the greatest trait diversity found among passerines. Among these species, telomeres are significantly shorter in fast-lived than in slow-lived species, suggesting that telomere length may have evolved to mediate trade-offs between physiological requirements underlying the diversity of pace-of-life strategies in birds. This association was attenuated when excluding studies that may include interstitial telomeres in the estimation of mean telomere length. Curiously, within some species, larger individual chromosome size predicts longer telomere lengths on that chromosome, leading to the hypothesis that telomere length also covaries with chromosome length across species. We show that longer mean chromosome length or genome size tends to be associated with longer mean early-life telomere length (measured across all chromosomes) within a phylogenetic framework constituting up to 31 bird species. These associations were strengthened when excluding highly influential outliers. However, sensitivity analyses suggested that they were susceptible to sample size effects and not robust to the exclusion of studies that may include interstitial telomeres. Combined, our analyses generalize patterns previously found within a few species and provide potential adaptive explanations for the 10-fold variation in telomere lengths observed among birds.

A trait-based approach to determining principles of plant biogeography.

Lineage-specific traits determine how plants interact with their surrounding environment. Unrelated species may evolve similar phenotypic characteristics to tolerate, persist in, and invade environments with certain characteristics, resulting in some traits becoming relatively more common in certain types of habitats. Analyses of these general patterns of geographical trait distribution have led to the proposal of general principles to explain how plants diversify in space over time. Trait-environment correlation analyses quantify to what extent unrelated lineages have similar evolutionary responses to a given type of habitat. In this synthesis, I give a short historical overview on trait-environment correlation analyses, from some key observations from classic naturalists to modern approaches using trait evolution models, large phylogenies, and massive data sets of traits and distributions. I discuss some limitations of modern approaches, including the need for more realistic models, the lack of data from tropical areas, and the necessary focus on trait scoring that goes beyond macromorphology. Overcoming these limitations will allow the field to explore new questions related to trait lability and niche evolution and to better identify generalities and exceptions in how plants diversify in space over time.

Evolution of female song and duetting in the chaffinch (Fringilla) species complex

Female song is ancestral to songbirds and shows considerable phylogenetic signal, but its presence also appears to be labile and correlated with life‐history and ecology. While previous studies have examined the evolution of female song across species‐rich families, here we studied female song in island populations of a recently diverged species‐complex, the chaffinches (genus Fringilla). We show this behaviour has recently evolved in these populations, and probably on two independent occasions. In F. teydea, female song is performed in loose duets with males, while in F. coelebs, female song is produced solo. Populations with singing females showed year‐round territoriality and were found in regions with low seasonality – both factors previously connected with high rates of female singing. To determine the relative saliency of female songs to conspecifics, and the degree to which they instigate territorial defence behaviours, we performed a series of speaker playback experiments. In F. c. canariensis, female song could induce comparable responses to male song, whilst duetting F. teydea playback produced similar responses to solo songs, and therefore may relate to within‐pair communication instead. Our results suggest female song can be a highly labile trait that can evolve over short evolutionary timescales.

Convergent evolution of the annual life history syndrome from perennial ancestors.

Despite most angiosperms being perennial, once-flowering annuals have evolved multiple times independently, making life history traits among the most labile trait syndromes in flowering plants. Much research has focused on discerning the adaptive forces driving the evolution of annual species, and in pinpointing traits that distinguish them from perennials. By contrast, little is known about how 'annual traits' evolve, and whether the same traits and genes have evolved in parallel to affect independent origins of the annual syndrome. Here, we review what is known about the distribution of annuals in both phylogenetic and environmental space and assess the evidence for parallel evolution of annuality through similar physiological, developmental, and/or genetic mechanisms. We then use temperate grasses as a case study for modeling the evolution of annuality and suggest future directions for understanding annual-perennial transitions in other groups of plants. Understanding how convergent life history traits evolve can help predict species responses to climate change and allows transfer of knowledge between model and agriculturally important species.

Nodule INception-independent epidermal events lead to bacterial entry during nodule development in peanut (Arachis hypogaea).

Legumes can host nitrogen-fixing rhizobia inside root nodules. In model legumes, rhizobia enter via infection threads (ITs) and develop nodules in which the infection zone contains a mixture of infected and uninfected cells. Peanut (Arachis hypogaea) diversified from model legumes c. 50-55 million years ago. Rhizobia enter through 'cracks' to form nodules in peanut roots where cells of the infection zone are uniformly infected. Phylogenomic studies have indicated symbiosis as a labile trait in peanut. These atypical features prompted us to investigate the molecular mechanism of peanut nodule development. Combining cell biology, genetics and genomic tools, we visualized the status of hormonal signaling in peanut nodule primordia. Moreover, we dissected the signaling modules of Nodule INception (NIN), a master regulator of both epidermal infection and cortical organogenesis. Cytokinin signaling operates in a broad zone, from the epidermis to the pericycle inside nodule primordia, while auxin signaling is narrower and focused. Nodule INception is involved in nodule organogenesis, but not in crack entry. Nodulation Pectate Lyase, which remodels cell walls during IT formation, is not required. By contrast, Nodule enhanced Glycosyl Hydrolases (AhNGHs) are recruited for cell wall modification during crack entry. While hormonal regulation is conserved, the function of the NIN signaling modules is diversified in peanut.

Diversification and trait evolution in New Zealand woody lineages across changing biomes

ABSTRACT Diversification of woody plant lineages in New Zealand has unfolded in complex physiographic, climatic, and environmental contexts. Many tree and shrub lineages have existed in New Zealand since the late Cenozoic when Forest was the dominant biome, subsequently diversifying (or continuing to diversify) during the Pliocene/Pleistocene as Open (below treeline) and Alpine biomes emerged. We examine the links between biomes occupied, traits, and diversification. In particular, whether traits are phylogenetically conserved or ecologically constrained and their relationship to biomes occupied. We focus on Melicytus, Myrsine and Pseudopanax which occur across Forest, Open, and Alpine biomes. Our approach combines measured traits and modelled niche traits of extant species to examine the importance of biome occupancy and biome shifts on trait evolution in these lineages. Our results demonstrate trait values are filtered by biomes in these lineages and can predict biomes occupied. However, few biome shifts were associated with trait evolution, typically only biome shifts into extreme environments (Alpine) involved trait innovations. In addition to biomes, trait evolution can also be influenced by species age, trait lability and broad climatic change. Integrating functional traits in a phylogenetic framework can identify how evolutionary and ecological features create modern biogeographic patterns in New Zealand.

Effects of laboratory holding time and diet type on labile traits in the crab Cancer irroratus Say, 1817 (Decapoda: Brachyura: Cancridae)

Abstract Collecting animals from the field and bringing them into the laboratory elicits acute and chronic stress responses that may affect the interpretation of experimental outcomes. The effects of prolonged laboratory holding (three months) on labile traits (metabolic rate and spontaneous activity) were quantified for the Atlantic rock crab Cancer irroratusSay, 1817. The effects of diet (heterogenous versus homogenous) on crab condition (hemolymph protein density, crab mass, and chelal compression strength) were also assessed. When offered a heterogeneous diet C. irroratus displayed a clear preference for mussels and an aversion to herring and algae. The amount crabs ate in the laboratory was negatively correlated to the density of hemolymph protein at the time of collection, which affirms the strong link between nutritional status and hemolymph protein in crustaceans. It also suggests that crabs in good nutritional condition may forgo eating even a high-quality meal if they are stressed. Overall, providing a heterogenous diet did not significantly improve survival rates or affect labile-trait responses in crabs. In contrast, prolonged holding in the laboratory had considerable effects on labile traits: resting metabolic rate (RMR) was highest after one week, but declined over the holding period. There was also a reduction in variation of locomotor activity for approximately 4 weeks. Acute stress responses (increased RMR and activity) also occurred after transfer from holding tanks to experimental chambers, likely due to animal handling. Given the increasing attention paid to animal sentience and welfare, especially for decapod crustaceans, the amount of time that wild crustaceans are held in the laboratory should be considered carefully.

Structural analysis of extrafloral nectaries of Senna occidentalis L.: insights on diversity and evolution.

The extrafloral nectaries of S. occidentalis were studied structurally and anatomically (at secretory and post-secretory developmental stages). Role of extrafloral nectaries as a common plant-adoptive characteristic in context to diversity and phylogenetic pattern was also speculated while exploring other collaborative evolutionary implications of this plant. Extrafloral nectaries (EFNs) are widespread and evolutionarily labile traits that have repeatedly and remarkably evolved in vascular plants. Morphological descriptions of the EFNs of certain plant species are common in the literature, but they rarely relate morphology with histology, gland distribution and secretory characteristics. Studies relating EFNs features, i.e., morphology and distribution with their differential visitation by insects, viz. ants and the cost of maintenance to the plants are important to understand the evolution of these glands. Therefore, in this study a morphological, anatomical (structure and ultrastructure) and secretory characterization of EFNs occurring on Senna occidentalis L. is made with the implications of gland attributes discussed from a functional perspective. S. occidentalis L. (Caesalpiniaceae) is an economically important species from industrial, medicinal and agricultural perspective. Observations from the result showed that shape of the EFNs (size 1-2mm) ranged to globular, ovoid-conical, dome-shaped, fusiform or cylindrical with conical tip. The EFNs were sessile, positioned interpetiolar or seated at the base of petiole. Light and transmission electron microscopic studies showed the specific internal structures of the extrafloral nectary. Two developmental stages of the EFNs (secretory and post-secretory) were recognized. Our current understanding of the phylogenetic patterns of EFNs makes them powerful candidates for future work exploring the drivers of their evolutionary origins, shifts, and losses.

Unifying individual differences in personality, predictability and plasticity: A practical guide

Abstract Organisms use labile traits to respond to different conditions over short time‐scales. When a population experiences the same conditions, we might expect all individuals to adjust their trait expression to the same, optimal, value, thereby minimising phenotypic variation. Instead, variation abounds. Individuals substantially differ not only from each other, but also from their former selves, with the expression of labile traits varying both predictably and unpredictably over time. A powerful tool for studying the evolution of phenotypic variation in labile traits is the mixed model. Here, we review how mixed models are used to quantify individual differences in both means and variability, and their between‐individual correlations. Individuals can differ in their average phenotypes (e.g. behavioural personalities), their variability (known as ‘predictability’ or intra‐individual variability), and their plastic response to different contexts. We provide detailed descriptions and resources for simultaneously modelling individual differences in averages, plasticity and predictability. Empiricists can use these methods to quantify how traits covary across individuals and test theoretical ideas about phenotypic integration. These methods can be extended to incorporate plastic changes in predictability (termed ‘stochastic malleability’). Overall, we showcase the unfulfilled potential of existing statistical tools to test more holistic and nuanced questions about the evolution, function, and maintenance of phenotypic variation, for any trait that is repeatedly expressed.

Phytochemistry reflects different evolutionary history in traditional classes versus specialized structural motifs

Foundational hypotheses addressing plant–insect codiversification and plant defense theory typically assume a macroevolutionary pattern whereby closely related plants have similar chemical profiles. However, numerous studies have documented variation in the degree of phytochemical trait lability, raising the possibility that phytochemical evolution is more nuanced than initially assumed. We utilize proton nuclear magnetic resonance (1H NMR) data, chemical classification, and double digest restriction-site associated DNA sequencing (ddRADseq) to resolve evolutionary relationships and characterize the evolution of secondary chemistry in the Neotropical plant clade Radula (Piper; Piperaceae). Sequencing data substantially improved phylogenetic resolution relative to past studies, and spectroscopic characterization revealed the presence of 35 metabolite classes. Metabolite classes displayed phylogenetic signal, whereas the crude 1H NMR spectra featured little evidence of phylogenetic signal in multivariate tests of chemical resonances. Evolutionary correlations were detected in two pairs of compound classes (flavonoids with chalcones; p-alkenyl phenols with kavalactones), where the gain or loss of a class was dependent on the other’s state. Overall, the evolution of secondary chemistry in Radula is characterized by strong phylogenetic signal of traditional compound classes and weak phylogenetic signal of specialized chemical motifs, consistent with both classic evolutionary hypotheses and recent examinations of phytochemical evolution in young lineages.

Behavioural phenotype modulates group size effects in a lizard

Behaviour is a highly labile trait that can be rapidly modified to mitigate the effects of changing environmental conditions. Among the biotic and abiotic factors acting to prompt plastic responses, the social environment has been proposed as being one of the primary modulating forces on behaviour. Being part of a group has particular influence on the expression of risky behaviour in that added eyes and ears serve to decrease a group member's vulnerability to predation resulting in the mean behavioural expression of behaviours such as activity and/or exploration increasing with group size. A large body of work has documented such group size effects. However, as this process may operate at the individual level it is unclear how the social environment affects the consistent expression of personality. Thus, we examined the interactive effects of behavioural phenotype and social context on the stability of individual behaviour in the delicate skink, Lampropholis delicata . Lizards were exposed to a series of assays measuring activity, exploration and boldness in groups of one, two, four and eight. Repeatability was determined across group size treatments and the combined effects of social context and behavioural type on behavioural plasticity were assessed. We found that the predicted patterns of group size effects were only observed when each lizard's behavioural phenotype was considered. Inactive and shy lizards increased their behaviour with increasing group size whereas active and bold lizards exhibited the opposite pattern. Additionally, the degree to which an individual adjusted its behavioural response was contingent upon its behavioural phenotype, with slow lizards showing higher levels of responsiveness than fast lizards. Despite this plasticity, between-individual differences in the expression of activity, exploration and boldness persisted. Thus, our study provides strong evidence that the effects of an individual's personality are stronger than those of group size. • Predicted group size effects are evident only when behavioural type is considered. • Social context influences the expression of consistent individual behaviour. • Personality dictates degree and direction of responses towards the social context.

Diversification, disparification and hybridization in the desert shrubs Encelia.

There are multiple hypotheses for the spectacular plant diversity found in deserts. We explore how different factors, including the roles of ecological opportunity and selection, promote diversification and disparification in Encelia, a lineage of woody plants in the deserts of the Americas. Using a nearly complete species-level phylogeny based on double-digest restriction-aided sequencing along with a broad set of phenotypic traits, we estimate divergence times and diversification rates, identify instances of hybridization, quantify trait disparity and assess phenotypic divergence across environmental gradients. We show that Encelia originated and diversified recently (mid-Pleistocene) and rapidly, with rates comparable to notable adaptive radiations in plants. Encelia probably originated in the hot deserts of North America, with subsequent diversification across steep environmental gradients. We uncover multiple instances of gene flow between species. The radiation of Encelia is characterized by fast rates of phenotypic evolution, trait lability and extreme disparity across environments and between species pairs with overlapping geographic ranges. Encelia exemplifies how interspecific gene flow in combination with high trait lability can enable exceptionally fast diversification and disparification across steep environmental gradients.

Evolution of stamen dimetrism in Melastomataceae, a large radiation of pollen flowers

Changes in floral traits across evolutionary time are expected in response to selective pressures imposed by pollinators. Stamen dimetrism (here defined as size differences between stamens within the same flower) represents an important strategy to decrease pollen loss during bee-flower interactions in pollen flowers. However, the evolutionary history of stamen dimetrism, and the links between this and other reproductive traits across long periods of time are still poorly understood. Here we investigate the evolution of stamens dimetrism and the evolutionary correlation of this trait and other floral structures and reproductive strategies in Melastomataceae. Floral traits were scored from 336 species and reproductive biology data was gathered for 81 species. Stamen dimetrism is a labile trait, appeared several times throughout the evolutionary history of this clade and is evolutionarily correlated to floral size. Among the 81 species analyzed, we observed that the lineages that depend on pollinators to reproduce correspond to those that evolved the highest stamen dimetrism. The evolutionary lability of stamen dimetrism has probably contributed to the maintenance of the buzz pollination adaptive plateau in possibly the largest radiation of pollen flowers in angiosperms.