Papers by Carlos Murga Herrera
SummaryEpigenetic mosaicism is a possible source of within-plant phenotypic heterogeneity, yet it... more SummaryEpigenetic mosaicism is a possible source of within-plant phenotypic heterogeneity, yet its frequency and developmental origin remain unexplored. This study examines whether the extant epigenetic heterogeneity within long-lived Lavandula latifolia (Lamiaceae) shrubs reflects recent epigenetic modifications experienced independently by different plant parts or, alternatively, it is the cumulative outcome of a steady lifetime process.Leaf samples from different architectural modules were collected from three L. latifolia plants and characterized epigenetically by global DNA cytosine methylation and methylation state of methylation-sensitive amplified fragment length polymorphism markers (MS-AFLP). Epigenetic characteristics of modules were then assembled with information on the branching history of plants. Methods borrowed from phylogenetic research were used to assess genealogical signal of extant epigenetic variation and reconstruct within-plant genealogical trajectory of epi...
Pollinators can mediate facilitative or competitive relationships between plant species, but the ... more Pollinators can mediate facilitative or competitive relationships between plant species, but the comparative importance of these two conflicting phenomena in shaping community-wide pollinator resource use remains unexplored. This paper examines the idea that the arrangement in pollinator niche space of plant species samples comprising complete or nearly complete regional or local plant communities can help to evaluate the relative importance of facilitation and competition as drivers of community-wide pollinator resource use. Pollinator composition data for insect-pollinated plants from the Sierra de Cazorla mountains (southeastern Spain), comprising 85% of families and ~95% of widely distributed insect-pollinated species, were used to address the following questions at regional (45 sites, 221 plant species) and local (one site, 73 plant species) spatial scales: (1) Do objectively identifiable plant species clusters occur in pollinator niche space ? Four different pollinator niche s...
Ecological Monographs, 2019
Pollinator service is essential for successful sexual reproduction and long-term population persi... more Pollinator service is essential for successful sexual reproduction and long-term population persistence of animal-pollinated plants, and innumerable studies have shown that insufficient service by pollinators results in impaired sexual reproduction ("pollen limitation"). Studies directly addressing the predictors of variation in pollinator service across species or habitats remain comparatively scarce, which limits our understanding of the primary causes of natural variation in pollen limitation. This paper evaluates the importance of pollinationrelated features, evolutionary history, and environment as predictors of pollinator service in a large sample of plant species from undisturbed montane habitats in southeastern Spain. Quantitative data on pollinator visitation were obtained for 191 insect-pollinated species belonging to 142 genera in 43 families, and the predictive values of simple floral traits (perianth type, class of pollinator visitation unit, and visitation unit dry mass), phylogeny, and habitat type were assessed. A total of 24,866 pollinator censuses accounting for 5,414,856 flower-minutes of observation were conducted on 510 different dates. Flowering patch and single flower visitation probabilities by all pollinators combined were significantly predicted by the combined effects of perianth type (open vs. restricted), class of visitation unit (single flower vs. flower packet), mass of visitation unit, phylogenetic relationships, and habitat type. Pollinator composition at insect order level varied extensively among plant species, largely reflecting the contrasting visitation responses of Coleoptera, Diptera, Hymenoptera, and Lepidoptera to variation in floral traits. Pollinator composition had a strong phylogenetic component, and the distribution of phylogenetic autocorrelation hotspots of visitation rates across the plant phylogeny differed widely among insect orders. Habitat type was a key predictor of pollinator composition, as major insect orders exhibited decoupled variation across habitat types in visitation rates. Comprehensive pollinator sampling of a regional plant community has shown that pollinator visitation and composition can be parsimoniously predicted by a combination of simple floral features, habitat type, and evolutionary history. Ambitious community-level studies can help to formulate novel hypotheses and questions, shed fresh light on long-standing controversies in pollination research (e.g., "pollination syndromes"), and identify methodological cautions that should be considered in pollination community studies dealing with small, phylogenetically biased plant species samples.
Evidence for pollinator declines largely originates from mid-latitude regions in North America an... more Evidence for pollinator declines largely originates from mid-latitude regions in North America and Europe. Geographical heterogeneity in pollinator trends combined with geographical biases in pollinator studies, can produce distorted extrapolations and limit understanding of pollinator responses to environmental changes. In contrast to the declines experienced in some well-investigated European and North American regions, honeybees seem to have increased recently in some areas of the Mediterranean Basin. Since honeybees can impact negatively on wild bees, it was hypothesized that a biome-wide alteration in bee pollinator assemblages may be underway in the Mediterranean Basin involving a reduction in the importance of wild bees as pollinators. This hypothesis was tested using published quantitative data on bee pollinators of wild and cultivated plants from studies conducted between 1963-2017 in 13 circum-Mediterranean countries. Honeybee colonies increased exponentially and wild bees...
American Journal of Botany, 2019
Phenotypic heterogeneity of reiterated, homologous structures produced by individual plants has e... more Phenotypic heterogeneity of reiterated, homologous structures produced by individual plants has ecological consequences for plants and their animal consumers. This paper examines experimentally the epigenetic mosaicism hypothesis, which postulates that within-plant variation in traits of reiterated structures may partly arise from different parts of the same genetic individual differing in patterns or extent of genomic DNA methylation. METHODS: Leaves of paired ramets borne by field-growing Helleborus foetidus plants were infiltrated periodically over the entire flowering period with either a water solution of the demethylating agent zebularine or just water as the control. The effects of the zebularine treatment were assessed by quantifying genome-wide DNA cytosine methylation in leaves and monitoring inflorescence growth and flower production, number of ovules per flower, pollination success, fruit set, seed set, seed size, and distribution of sap-feeding insects. RESULTS: Genomic DNA from leaves in zebularine-treated ramets was significantly less methylated than DNA from leaves in control ones. Inflorescences in treated ramets grew smaller and produced fewer flowers, with fewer ovules and lower follicle and seed set, but did not differ from inflorescences in untreated ramets in pollination success or seed size. The zebularine treatment influenced the within-plant distribution of sap-feeding insects. CONCLUSIONS: Experimental manipulation of genomic DNA methylation level in leaves of wild-growing H. foetidus plants induced considerable within-plant heterogeneity in phenotypic (inflorescences, flowers, fecundity) and ecologically relevant traits (herbivore distribution), which supports the hypothesis that epigenetic mosaicism may partly account for within-plant variation.
Web Ecology, 2017
Diversification of ecology into subdisciplines that run from macroecology to landscape, community... more Diversification of ecology into subdisciplines that run from macroecology to landscape, community, and population ecology largely reflects its specialization on different segments of the spatial gradient over which recognizable ecological patterns and processes occur. In all these cases, the elemental units involved in the patterns and processes of interest to ecologists are individuals from the same or different species. No distinct flavor of ecology has yet emerged that focuses on patterns and processes revolving around the lowermost end of the spatial gradient, which in the case of plants corresponds to the within-individual domain. Intraindividual heterogeneity in organ traits, however, is quantitatively important and has multiple consequences for plant individuals, populations, and communities, and for animal consumers as well. This paper first provides an overview of current knowledge on plant traits that vary subindividually, the magnitude of subindividual variation, and its spatial patterning. Examples will then be presented on the consequences of subindividual variation for plants and animal consumers at individual, population, or community levels. Finally, the recently emerging links between genetics, epigenetics, subindividual variation, and population ecology will be illustrated using results on variation in seed size, a functional plant trait playing an important role in plant population dynamics. Further observational and experimental studies are needed which link ecological and phenotypic measurements of plants to their epigenetic and genetic characteristics, in order to understand the three-way relationships between subindividual variability, genetic features, and epigenetic mosaicism. Another proposed line of inquiry should focus on evaluating whether subindividual epigenetic mosaics eventually translate into epigenetically heterogeneous progeny, thus contributing to the maintenance of population and community functional diversity.
Epigenomes, 2017
Experimental alteration of DNA methylation is a suitable tool to infer the relationship between p... more Experimental alteration of DNA methylation is a suitable tool to infer the relationship between phenotypic and epigenetic variation in plants. A detailed analysis of the genome-wide effect of demethylating agents, such as 5-azacytidine (5azaC), and zebularine is only available for the model species Arabidopsis thaliana, which suggests that 5azaC may have a slightly larger effect. In this study, global methylation estimates obtained by high-performance liquid chromatography (HPLC) analyses were conducted to investigate the impact of 5azaC treatment on leaf and root tissue in Erodium cicutarium (Geraniaceae), which is an annual herb native to Mediterranean Europe that is currently naturalized in all continents, sometimes becoming invasive. We used seeds collected from two natural populations in SE Spain. Root tissue of the second generation (F2) greenhouse-grown seedlings had a significantly lower global cytosine methylation content than leaf tissue (13.0 vs. 17.7% of all cytosines). Leaf tissue consistently decreased methylation after treatment, but the response of root tissue varied according to seed provenance, suggesting that genetic background can mediate the response to experimental demethylation. We also found that both leaf number and leaf length were reduced in treated seedlings supporting a consistent phenotypic effect of the treatment regardless of seedling provenance. These findings suggest that, although the consequences of experimental demethylation may be tissue-and background-specific, this method is effective in altering early seedling development, and can thus be useful in ecological epigenetic studies that are aiming to investigate the links between epigenetic and phenotypic variation in non-model plant species.
Ecological Monographs, 2018
Current notions of "pollinator decline" and "pollination crisis" mainly arose from studies on pol... more Current notions of "pollinator decline" and "pollination crisis" mainly arose from studies on pollinators of economic value in anthropogenic ecosystems of mid-latitude temperate regions. Comprehensive long-term pollinator data from biologically diverse, undisturbed communities are needed to evaluate the actual extent of the so-called "global pollination crisis." This paper analyzes the long-term dynamics of pollinator abundance in undisturbed Mediterranean montane habitats using pollinator visitation data for 65 plant species collected over two decades. Objectives are (1) to elucidate patterns of long-term changes in pollinator abundance from the perspectives of individual plant species, major pollinator groups, and the whole plant community and (2) to propose a novel methodological implementation based on combining a planned missing data design with the analytical strength of mixed effects models, which allows one to draw community-wide inferences on long-term pollinator trends in species-rich natural habitats. Probabilistic measurements ("patch visitation probability" and "flower visitation probability" per time unit) were used to assess pollinator functional abundance for each plant species on two separate, randomly chosen years. A total of 13,054 pollinator censuses accounting for a total watching effort of 2,877,039 flower-min were carried out on 299 different dates. Supra-annual unstability in pollinator functional abundance was the rule, with visitation probability to flowering patches and/or individual flowers exhibiting significant heterogeneity between years in the majority of plant species (83%). At the plant-community level, there was a significant linear increase in pollinator functional abundance over the study period. Probability of pollinator visitation to flowering patches and individual flowers increased due to increasing visitation by small solitary bees and, to a lesser extent, small beetles. Visitation to different plant species exhibited contrasting changes, and insect orders and genera differed widely in sign and magnitude of linear abundance trends, thus exemplifying the complex dynamics of community-wide changes in pollinator functional abundance. Results of this investigation indicate that pollinator declines are not universal beyond anthropogenic ecosystems; stress the need for considering broader ecological scenarios and comprehensive samples of plants and pollinators; and illustrate the crucial importance of combining ambitious sampling designs with powerful analytical schemes to draw reliable inferences on pollinator trends at the plant community level.
American journal of botany, Apr 4, 2018
The ecological and evolutionary significance of natural epigenetic variation (i.e., not based on ... more The ecological and evolutionary significance of natural epigenetic variation (i.e., not based on DNA sequence variants) variation will depend critically on whether epigenetic states are transmitted from parents to offspring, but little is known on epigenetic inheritance in nonmodel plants. We present a quantitative analysis of transgenerational transmission of global DNA cytosine methylation (= proportion of all genomic cytosines that are methylated) and individual epigenetic markers (= methylation status of anonymous MSAP markers) in the shrub Lavandula latifolia. Methods based on parent-offspring correlations and parental variance component estimation were applied to epigenetic features of field-growing plants ('maternal parents') and greenhouse-grown progenies. Transmission of genetic markers (AFLP) was also assessed for reference. Maternal parents differed significantly in global DNA cytosine methylation (range = 21.7-36.7%). Greenhouse-grown maternal families differed s...
American journal of botany, Jan 16, 2017
Epigenetic variation can play a role in local adaptation; thus, there should be associations amon... more Epigenetic variation can play a role in local adaptation; thus, there should be associations among epigenetic variation, environmental variation, and functional trait variation across populations. This study examines these relationships in the perennial herb Helleborus foetidus (Ranunculaceae). Plants from 10 subpopulations were characterized genetically (AFLP, SSR markers), epigenetically (MSAP markers), and phenotypically (20 functional traits). Habitats were characterized using six environmental variables. Isolation-by-distance (IBD) and isolation-by-environment (IBE) patterns of genetic and epigenetic divergence were assessed, as was the comparative explanatory value of geographical and environmental distance as predictors of epigenetic, genetic, and functional differentiation. Subpopulations were differentiated genetically, epigenetically, and phenotypically. Genetic differentiation was best explained by geographical distance, while epigenetic differentiation was best explained...
Flora, 2017
In habitats where low ambient temperature limits entomophilous pollination of early-blooming plan... more In habitats where low ambient temperature limits entomophilous pollination of early-blooming plants, intrafloral warming facilitates pollinator visitation and enhances plant reproductive success. A novel mechanism for intrafloral warming was described recently for the early-blooming, bumblebee pollinated herb Helleborus foetidus (Ranunculaceae) where the warming agent was the metabolic heat dissipated by dense populations of the nectar-specialist yeast Metschnikowia reukaufii. This paper reports results of field experiments conducted over three consecutive years aimed at (1) ascertaining whether flowers of H. foetidus with warmed interiors differed in pollination success from flowers experiencing natural intrafloral microclimate; and (2) testing the hypothesis that the pollination effects of intrafloral warming should be stronger during the early stages of the H. foetidus flowering season, when inclement weather and low ambient temperatures often limit pollinator activity. Intrafloral warming resulted in increased number of pollen tubes per style and reduced probability of pollen limitation early in the flowering season, while these effects were reversed in late-season flowers. Since the number of pollen tubes was directly correlated with number of pollen grains on the stigma, fruit set and number of seeds produced by individual flowers, results of this study are interpreted as indicating that yeast-induced intrafloral warming has the potential to influence reproductive success in H. foetidus via effects on pollinator activity, although the nature of effects will vary depending on environmental conditions. Results also stress the importance of ambient and floral temperature for the pollination of early-blooming plants.
Ecology and Evolution, 2016
Little is known on the potential of ecological disturbance to cause genetic and epigenetic change... more Little is known on the potential of ecological disturbance to cause genetic and epigenetic changes in plant populations. We take advantage of a long-term field experiment initiated in 1986 to study the demography of the shrub Lavandula latifolia, and compare genetic and epigenetic characteristics of plants in two adjacent subplots, one experimentally disturbed and one left undisturbed, 20 years after disturbance. Experimental setup was comparable to an unreplicated 'Before-After-Control-Impact' (BACI) design where a single pair of perturbed and control areas were compared. When sampled in 2005, plants in the two subplots had roughly similar ages, but they had established in contrasting environments: dense conspecific population ('Undisturbed' subpopulation) versus open area with all conspecifics removed ('Disturbed' subpopulation). Plants were characterized genetically and epigenetically using amplified fragment length polymorphism (AFLP) and two classes of methylation-sensitive AFLP (MSAP) markers. Subpopulations were similar in genetic diversity but differed in epigenetic diversity and multilocus genetic and epigenetic characteristics. Epigenetic divergence between subpopulations was statistically unrelated to genetic divergence. Bayesian clustering revealed an abrupt linear boundary between subpopulations closely coincident with the arbitrary demarcation line between subplots drawn 20 years back, which supports that genetic and epigenetic divergence between subpopulations was caused by artificial disturbance. There was significant fine-scale spatial structuring of MSAP markers in both subpopulations, which in the Undisturbed one was indistinguishable from that of AFLP markers. Genetic differences between subpopulations could be explained by divergent selection alone, while the concerted action of divergent selection and disturbance-driven appearance of new methylation variants in the Disturbed subpopulation is proposed to explain epigenetic differences. This study provides the first empirical evidence to date suggesting that relatively mild disturbances could leave genetic and epigenetic signatures on the next adult generation of long-lived plants.
Molecular ecology, Apr 6, 2016
Despite the recent upsurge of interest on natural epigenetic variation of non-model organisms, fa... more Despite the recent upsurge of interest on natural epigenetic variation of non-model organisms, factors conditioning the spatial structure of epigenetic diversity in wild plant populations remain virtually unexplored. We propose that information on processes shaping natural epigenetic variation can be gained by using the spatial structure of genetic diversity as null model. Departures of epigenetic isolation-by-distance (IBD) patterns from genetic IBD patterns for the same sample, particularly differences in slope of similarity-distance regressions, will reflect the action of factors that operate specifically on epigenetic variation, including imperfect transgenerational inheritance and responsiveness to environmental factors of epigenetic marks. As a proof of concept, we provide a comparative analysis of spatial genetic and epigenetic structure of 200 mapped individuals of the perennial herb Helleborus foetidus. Plants were fingerprinted using nuclear microsatellites, amplified frag...
Anales del Jardín Botánico de Madrid, 2015
Comparative effects of two species of floricolous Metschnikowia yeasts on nectar. Anales Jard. Bo... more Comparative effects of two species of floricolous Metschnikowia yeasts on nectar. Anales Jard. Bot. Madrid 72(1): e019 Nectar yeast communities in southern Spain are dominated by two closely-related species, Metschnikowia reukaufii Pitt & M.W. Mill. and M. gruessii Gim.-Jurado (Ascomycota, Saccharomycetales), although they tend to be distributed differentially across different host plants. We explore here the possibility that the two yeasts play different functional roles in floral nectar by differing in their impact on sugar concentration and composition of nectar. Experiments were undertaken under controlled conditions using bumblebees caught foraging on the flowers of two different host plants each of which is known to harbor predominantly one of the two yeasts. Bumblebees were used as sources of inocula to obtain two groups of samples from the nectar of Helleborus foetidus L. (Ranunculaceae): nectar samples inoculated with M. gruessii and samples inoculated with M. reukaufii. Metschnikowia gruessii was poorly represented in nectar samples, while M. reukaufii was by far the most common and had the highest cell density. Although the two yeasts caused relatively similar changes in nectar sugar composition, which involved increasing fructose and decreasing sucrose proportions, they marginally differed in their quantitative impact on total nectar sugar concentration. Results suggest that differential yeast occurrence across host plants may lead to yeast specialization and modify the outcomes of the plant-pollinator interface.
Annals of botany, 2010
Persistence of withered corollas after anthesis ('corolla marcescence') is widespread in ... more Persistence of withered corollas after anthesis ('corolla marcescence') is widespread in angiosperms, yet its functional significance does not seem to have been explored for any species. This note reports the results of experiments assessing the fecundity effects of marcescent corollas in two southern Spanish insect-pollinated plants, Lavandula latifolia (Lamiaceae) and Viola cazorlensis (Violaceae). The effect of marcescent corollas on seed production was evaluated experimentally on wild-growing plants. Newly open flowers were randomly assigned to either control or treatment groups in experimental plants. After anthesis, withered corollas of treatment flowers were removed and those in control flowers were left in place. Fruits produced by treatment and control flowers were collected shortly before dehiscence and the number of seeds counted. In V. cazorlensis, removal of withered corollas had no effect on percentage of fruit set, but mean seeds per fruit increased from 9·5 t...
Methods in molecular biology (Clifton, N.J.), 2012
Methods for estimating quantitative trait heritability in wild populations have been developed in... more Methods for estimating quantitative trait heritability in wild populations have been developed in recent years which take advantage of the increased availability of genetic markers to reconstruct pedigrees or estimate relatedness between individuals, but their application to real-world data is not exempt from difficulties. This chapter describes a recent marker-based technique which, by adopting a genomic scan approach and focusing on the relationship between phenotypes and genotypes at the individual level, avoids the problems inherent to marker-based estimators of relatedness. This method allows the quantification of the genetic component of phenotypic variance ("degree of genetic determination" or "heritability in the broad sense") in wild populations and is applicable whenever phenotypic trait values and multilocus data for a large number of genetic markers (e.g., amplified fragment length polymorphisms, AFLPs) are simultaneously available for a sample of ind...
American journal of botany, 2015
• Premise of the study: Continuous within-plant variation in quantitative traits of reiterated, h... more • Premise of the study: Continuous within-plant variation in quantitative traits of reiterated, homologous structures is a component of intraspecifi c variation, but its contribution to functional diversity remains largely unexplored. For the perennial Helleborus foetidus , we measured functional leaf traits to quantify the contribution of within-plant variation to intraspecifi c functional variance and evaluate whether within-plant variability itself deserves separate consideration. • Methods: Within-individual variation in eight leaf traits was quantifi ed for 138 plants sampled from 10 widely spaced locations in the Sierra de Cazorla, southeastern Spain. An amplifi ed fragment length polymorphism (AFLP) technique was used to look for associations between within-plant variability and specifi c AFLP markers. • Key results: Leafl ets from basal positions in ramets were longer, heavier, had greater surface area and larger stomata, and lower specifi c area, stomatal index, and stomatal density than those from distal positions. Continuous variation between leaves from the same ramet was the main source of population-wide variance for most traits. Within-plant variability differed among populations. Individuals differed in within-plant variability, which was largely independent of trait means and associated with genetic characteristics. Up to four AFLP markers were associated with the within-plant variability level of a given leaf trait. • Conclusions: Subindividual variability in continuous leaf traits was independent of plant means and related to genetic features. The within-individual component generally exceeded the between-individual component of intraspecifi c variance. Withinplant variation may broaden the ecological breadth and enhance stability and persistence of plant populations and communities and may provide novel insights when incorporated in trait-based community ecology models.
Molecular ecology, 2014
Inferences about the role of epigenetics in plant ecology and evolution are mostly based on studi... more Inferences about the role of epigenetics in plant ecology and evolution are mostly based on studies of cultivated or model plants conducted in artificial environments. Insights from natural populations, however, are essential to evaluate the possible consequences of epigenetic processes in biologically realistic scenarios with genetically and phenotypically heterogeneous populations. Here, we explore associations across individuals between DNA methylation transmissibility (proportion of methylation-sensitive loci whose methylation status persists unchanged after male gametogenesis), genetic characteristics (assessed with AFLP markers), seed size variability (within-plant seed mass variance), and realized maternal fecundity (number of recently recruited seedlings), in three populations of the perennial herb Helleborus foetidus along a natural ecological gradient in southeastern Spain. Plants (sporophytes) differed in the fidelity with which DNA methylation was transmitted to descenda...
Annals of Botany, 2012
† Background and Aims Variation in the composition of floral nectar reflects intrinsic plant char... more † Background and Aims Variation in the composition of floral nectar reflects intrinsic plant characteristics as well as the action of extrinsic factors. Microorganisms , particularly yeasts, represent one extrinsic factor that inhabit the nectar of animal-pollinated flowers worldwide. In this study a 'microbial imprint hypothesis' is formulated and tested, in which it is proposed that natural community-wide variation in nectar sugar composition will partly depend on the presence of yeasts in flowers. † Methods Occurrence and density of yeasts were studied microscopically in single-flower nectar samples of 22 animal-pollinated species from coastal xeric and sub-humid tropical habitats of the Yucatán Peninsula, Mexico. Nectar sugar concentration and composition were concurrently determined on the same samples using high-performance liquid chromatography (HPLC) methods. † Key Results Microscopical examination of nectar samples revealed the presence of yeasts in nearly all plant species (21 out of 22 species) and in about half of the samples examined (51. 8 % of total, all species combined). Plant species and individuals differed significantly in nectar sugar concentration and composition, and also in the incidence of nectar yeasts. After statistically controlling for differences between plant species and individuals, nectar yeasts still accounted for a significant fraction of community-wide variance in all nectar sugar parameters considered. Significant yeast × species interactions on sugar parameters revealed that plant species differed in the nectar sugar correlates of variation in yeast incidence. † Conclusions The results support the hypothesis that nectar yeasts impose a detectable imprint on communitywide variation in nectar sugar composition and concentration. Since nectar sugar features influence pollinator attraction and plant reproduction, future nectar studies should control for yeast presence and examine the extent to which microbial signatures on nectar characteristics ultimately have some influence on pollination services in plant communities.
Molecular Ecology, 2014
The ecological significance of epigenetic variation has been generally inferred from studies on m... more The ecological significance of epigenetic variation has been generally inferred from studies on model plants under artificial conditions, but the importance of epigenetic differences between individuals as a source of intraspecific diversity in natural plant populations remains essentially unknown. This paper investigates the relationship between epigenetic variation and functional plant diversity by conducting epigenetic (methylation-sensitive amplified fragment length polymorphisms, MSAP) and genetic (amplified fragment length polymorphisms, AFLP) marker-trait association analyses for 20 wholeplant, leaf and regenerative functional traits in a large sample of wild-growing plants of the perennial herb Helleborus foetidus from ten sampling sites in southeastern Spain. Plants differed widely in functional characteristics, and exhibited greater epigenetic than genetic diversity, as shown by percent polymorphism of MSAP fragments (92%) or markers (69%) greatly exceeding that for AFLP ones (41%). After controlling for genetic structuring and possible cryptic relatedness, every functional trait considered exhibited a significant association with at least one AFLP or MSAP marker. A total of 27 MSAP (13.0% of total) and 12 AFLP (4.4%) markers were involved in significant associations, which explained on average 8.2% and 8.0% of trait variance, respectively. Individual MSAP markers were Accepted Article This article is protected by copyright. All rights reserved. more likely to be associated with functional traits than AFLP markers. Between-site differences in multivariate functional diversity was directly related to variation in multilocus epigenetic diversity after multilocus genetic diversity was statistically accounted for. Results suggest that epigenetic variation can be an important source of intraspecific functional diversity in H. foetidus, possibly endowing this species with the capacity to exploit a broad range of ecological conditions despite its modest genetic diversity.
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Papers by Carlos Murga Herrera