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Herrera, C. M.

Publications and source records attributed to Herrera, C. M..

16 recordsLinked to original sources

Flower engineering by yeasts: reversal of nectar sugar concentration gradient within Darwin's syndrome inflorescences

Yeasts thriving in floral nectar have a proven ecological engineering capacity, as they modify the internal environment and the rewards of individual flowers in ways that influence how plants and pollinators interact. This floricentric perpective, however, produces only a partial view of the potential role of yeasts in plant-pollinator interactions, and little is known on their possible engineering effects on whole inflorescences. This study assessed yeast ability to shape features of the vertical inflorescences of Gladiolus illyricus (Iridaceae), a Darwins syndrome plant (acropetalous protandrous flowers pollinated by insects foraging from bottom to top). When yeasts were excluded from inflorescences a declining gradient in nectar sugar concentration built up from the oldest, female-stage flowers at bottom up to the youngest, male-stage ones at top. This plant-intrinsic gradient was reversed in inflorescences with yeasts, where sugar concentration increased from the female-stage flowers harboring dense yeasts population to male-stage ones with few yeasts. The two species of yeasts involved (Metschnikowi reukaufii and M. gruessii) differed in their effects. Results add support to the growing consensus that specialized nectar yeasts can obfuscate intrinsic patterns of intra and interspecific variation in flowering features of plants at the level of individual flower, inflorescence, population and plant community.

ecology↗

On commonness and rarity of insect pollinators: patterns of species abundance and diversity in a hyperdiverse Mediterranean assemblage

This paper presents a comprehensive analysis of patterns of species abundance and diversity in a hyperdiverse insect pollinator assemblage from Mediterranean montane habitats of southeastern Spain. Data on pollinator visitation to flowers of the community of entomophilous plants (288 species) were gathered over a 29-year period, and [~]95% of individual pollinators recorded were identified to species, totalling 46,401 individuals in 845 species. Shapes of species abundance distributions (SADs) were virtually identical at the regional (N = 56 sites) and local (one intensively studied site) scales, remained invariant over the study period, and were best predicted by the log-series distribution. Pollinator diversity estimates corresponding to the first three Hill numbers (Species richness, Shannon diversity and Simpson diversity; 0D, 1D and 2D, respectively) were obtained for each plant species x site x year combinations (N = 472 "sampling occasions"). Pollinator diversity measures varied widely among plant species; their frequency distributions were unimodal and strongly right-skewed; and interspecific variation was related to plant phylogeny, floral features and pollinator visitation rates. Pollinator diversity of individual plant species depended on habitat type, with dolomitic outcrops, rock cliffs and forest interior harboring the least diverse pollinator assemblages. Variation of 0D, 1D and 2D among habitats and years was largely decoupled, revealing a complex patterning of pollinator species richness and dominance. Estimated proportions of undetected pollinator diversity ("dark diversity") varied with insect order (lowest for Lepidoptera, highest for Diptera). The adoption of community ecology tools for assessing pollinator diversity improves our ability to elucidate pollinator responses to natural and anthropogenic environmental change, and permits hitherto unexplored questions in pollination ecology.

ecology↗

Somatic epigenetic mosaicism within Pistacia terebinthus (Anacardiaceae) trees leaves signature in reproductive cells

Propagation of epimutations over the branching topology of individuals often produces epigenetic mosaicism in long-lived plants, but whether such epigenetic mosaics can bear some evolutionary relevance remains largely unknown. We test here whether epigenetic reprogramming during gametogenesis completely erases intraplant epigenetic variation or, alternatively, some "signature" of the somatic mosaicism persists into reproductive cells. Intraplant variation in global cytosine methylation of leaf and pollen DNA samples was studied in five adult males of the dioecious tree Pistacia terebinthus (Anacardiaceae). Paired samples of fresh expanding leaves and pollen grains were taken from five different twigs in each tree, and percent cytosine methylation estimated using HPLC. Trees were internally heterogeneous in global cytosine methylation of both leaf and pollen. Within-tree variance accounted for 49.7% and 30.5% of population-wide variance in methylation for leaves and pollen, respectively. Tree means and within-tree variances in pollen methylation were directly correlated, respectively, with tree means and within-tree variances in leaf methylation. As a consequence of the epigenetic variability accumulated in somatic tissues over the individual plants lifetimes being only weakly eroded by gametogenesis, intraplant epigenetic heterogeneity became a natural source of epigenetic gametic diversity in the wild-growing population of P. terebinthus trees studied.

evolutionary biology↗

Flower yeasts obfuscate intrinsic individual variation in nectar sugar concentration in a bumble bee-pollinated plant

Nectar-dwelling yeasts modulate the ecology of interactions between flowers and pollinators. We evaluate here the hypothesis that floral nectar traits, which are a central element in most plant-pollinator relationships, can largely represent the extended fungal phenotypes of nectar-dwelling yeasts. The following specific question is addressed, Do local genotypes of the specialist nectar yeast Metschnikowia reukaufii have the ability to obfuscate intrinsic individual variation among Helleborus foetidus plants in nectar sugar concentration ? An array of paired plant-yeast genotypes mimicking a natural field situation was created in the laboratory by inoculating field-collected nectar from different plant individuals with distinct yeast genotypes following a factorial design. Chemical analyses of nectar sugars before and after exposure to yeasts were performed using ion-exchange high performance liquid chromatography. Plant individual, yeast genotype, and their interaction all had strong, significant effects on sucrose concentration of H. foetidus nectar. Yeasts abolished 79% of intrinsic variance among plants in nectar sucrose concentration and profoundly reshaped patterns of intrinsic among-plant variation. Our results support the hypothesis that among-plant variation in nectar sugar concentration found by pollinators in the field can sometimes reflect the extended phenotypes of nectar yeasts more closely than intrinsic differences among the plants themselves.

ecology↗

Flowers that cool themselves: thermal ecology of summer-blooming thistles in hot Mediterranean environments

Flower exposure to high temperature reduces the production, viability and performance of pollen, ovules and seeds, which in turn reduces individual fecundity and risks the survival of populations. Autonomous flower cooling could help to alleviate exposure of pollen and ovules to harmful temperatures, yet investigations on the thermal ecology of flowers in hot environments are still needed to evaluate the frequency, magnitude and ecological significance of floral thermoregulatory cooling. This paper reports a study on the thermal ecology of flower heads (=capitula) of 15 species of summer-blooming Asteraceae, tribe Cardueae, from hot-dry habitats in the southern Iberian Peninsula. Temperature inside (Tin) and outside (Tout) capitula were assessed under natural field conditions by two complementary sampling and measurement procedures, which provided information on the relationships between the two temperatures at the levels of individual capitula ("continuous recording") and local plant populations ("instantaneous measurements"). Baselines for the Tin-Tout relationship in absence of physiological activity were obtained by exposing dehydrated capitula to variable ambient temperatures. To assess whether the co-flowering capitula of summer-blooming Asteraceae collectively defined a distinct thermal layer, the vertical distribution of capitula relative to the ground was quantified. Bees visiting capitula were watched and the temperature of the air beside the visited capitulum was measured. Capitula of all species were exposed to high ambient temperatures during long periods, yet their interior was cooler than the air for most of the time, with temperature differentials ({Delta}T=Tin-Tout) quite often approaching, and sometimes exceeding -10{degrees}C. The relationship between Tin and Tout was best described by a composite of steep and shallow linear relationships separated by a breakpoint ({Psi}, interspecific range=25{degrees}C-35{degrees}C). Capitula were weakly or not thermoregulated for Tout<{Psi}, but switched to intense, thermoregulated cooling when Tout>{Psi}. Narrow vertical distribution of capitula above ground and similar cooling responses by all species produced a refrigerated floral layer where most bees foraged at Tout>{Psi} and presumably visited cooled capitula. Thermoregulatory cooling of capitula can benefit plant reproduction by reducing pollen and ovule exposure to high temperature, but also the populations of insect mutualists and antagonists through habitat improvement via a refrigerated flower layer. OPEN RESEARCH STATEMENTData, metadata and R script will be made available at figshare.

ecology↗

Plant phenotypes as trajectories: 38-year monitoring of shrub population reveals selection on individual lifetime fecundity schedule

Defining a phenotype is sometimes problematic in the case of modularly-built, nonunitary organisms with indeterminate growth such as plants. This paper presents a proof of concept of the evolutionary significance of considering lifetime trajectories of individual plants as a component of their phenotypes. Size and inflorescence production were monitored for the whole reproductive lifespans of N = 128 individuals of the Mediterranean woody shrub Lavandula latifolia (Lamiaceae) over a 38-year period to address the following questions: Did individuals vary in lifetime trajectories of size and fecundity?, and Were parameters describing individual trajectories significant predictors of cumulative lifetime reproduction? Individuals differed widely in lifetime fecundity and in every parameter describing lifetime fecundity schedule (age at first and last reproduction, longevity, and mean, variance, skewness and kurtosis of lifetime temporal distribution of inflorescence production). Phenotypic selection analysis revealed significant relationships between parameters of individual lifetime schedules and a surrogate of relative fitness of individuals (cumulative lifetime production of inflorescences divided by the average for all individuals). Significant selection gradients involved positive and negative directional selection, as well as instances of nonlinear selection, which showed that plants with certain shapes of lifetime fecundity schedules had fitness advantage over others. The notion of plant phenotypes as trajectories was strongly supported by the combined findings that individuals differed greatly in their "appearances" with regard to the way in which size and fecundity unfolded over lifetime, and that selection efficaciously "saw" such variation.

ecology↗

The Chlamydia effector IncE employs two short linear motifs to reprogram host vesicle trafficking

Chlamydia trachomatis, a leading cause of bacteria sexually transmitted infections, creates a specialized intracellular replicative niche by translocation and insertion of a diverse array of effectors (Incs) into the inclusion membrane. Here, we characterize IncE, a multi-functional Inc that encodes two non-overlapping short linear motifs (SLiMs) within its short cytosolic C-terminus. The proximal SLiM mimics an R-SNARE motif to recruit syntaxin (STX) 7 and 12-containing vesicles to the inclusion. The distal SLiM mimics the Sorting Nexin (SNX) 5 and 6 cargo binding site to recruit SNX6-containing vesicles to the inclusion. By simultaneously binding to two distinct vesicle classes, IncE reprograms host cell trafficking to promote the formation of a class of hybrid vesicles at the inclusion that are required for C. trachomatis intracellular development. Our work suggests that Incs may have evolved SLiMs to facilitate rapid evolution in a limited protein space to disrupt host cell processes.

microbiology↗

A dynamic epigenetic perspective on above and below-ground phenotypic responses to drought: insights from global DNA methylation in Erodium cicutarium

O_LIPremise of the study. Mounting evidence supports the view that the responses of plants to environmental stress are mediated by epigenetic factors, including DNA methylation. Understanding the relationships between DNA methylation, plant development and individual fitness under contrasting environments is key to uncover the potential impact of epigenetic regulation on plant adaptation. Experimental approaches that combine a controlled alteration of epigenetic features with exposure to some relevant stress factor can contribute to this end. C_LIO_LIMethods. We combined the experimental application of a demethylating agent (5-azacytidine) with recurrent drought, and recorded their effects on above- and below-ground phenotypic traits related to early development, phenology and fitness in Erodium cicutarium from two provenances. C_LIO_LIKey results. We found that 5-azacytidine significantly reduced DNA methylation in leaf and root tissues. Moreover, it slowed plant development, delayed flowering, and reduced the number of inflorescences produced, and such detrimental effects occurred independently of water regime. Recurrent drought reduced final above- and below-ground biomass and total inflorescence production, and such negative effects were unaffected by artificial changes in DNA methylation. Increased fruit and seed-set were the only adaptive responses to drought observed in E. cicutarium, together with an increased number of flowers per inflorescence recorded in water stressed plants previously treated with 5-azacytidine. C_LIO_LIConclusion. Epigenetic effects can desynchronize plant growth, flowering and senescence among individual plants in both favourable and adverse environments. Future studies should focus on understanding intraspecific variation in the ability to change plant methylome in response to stress. C_LI

ecology↗

Thermal biology of bee pollinators: diversity and community-level correlates

Community-wide assembly of plant-pollinator systems depends on an intricate combination of biotic and abiotic factors, including heterogeneity among pollinators in thermal biology and responses to abiotic factors. Studies on the thermal biology of pollinators have mostly considered only one or a few species of plants or pollinators at a time, and the possible driving role of the diversity in thermal biology of pollinator asemblages at the plant community level remains largely unexplored. More specifically, it is unknown whether diversity in the thermal biology of bees, a major pollinator group worldwide, contributes to the assembly and maintenance of diverse bee communities, broadens the spectrum of possibilities available to bee-pollinated plants, facilitate interspecific partitioning of ecological gradients across habitats, seasons and time of day, and/or enhance plant pollination success through complementarity effects. The objectives of this study were to assess the diversity in thermal biology of the bee assemblage that pollinates plants in a Mediterranean montane area, evaluate its taxonomic and phylogenetic underpinnings, and elucidate whether there existed seasonal, daily, between-habitat or floral visitation correlates of bee thermal biology which could contribute to partition ecological gradients among plant and bee species. Thermal biology parameters were obtained in the laboratory (K, intrinsic warming constant) and the field (thoracic and ambient temperature at foraging site, Tth and Tair) on individual bees of a diverse sample (N = 204 bee species) comprising most bee pollinators of the regional plant community. Species-specific thermal biology parameters were combined with quantitative field data on bee pollinators and flower visitation for the regional community of entomophilous plants (N = 292 plant species). Results revealed that the regional bee assemblage harbored considerable diversity in thermal biology features, that such diversity was mostly taxonomically, phylogenetically and body-size structured, and that the broad interspecific heterogeneity in thermal biology represented in the bee community as a whole eventually translated into daily, seasonal, among-habitat and flower visitation patterns at the plant community level. This lends support to the hypothesis that broad diversity in thermal biology of bees can act enhancing opportunities for bee coexistence, spatio-temporal partitioning of floral resources, and plant pollination success.

ecology↗

Assessing the links between pollinators and the genetic and epigenetic features of plant species with contrasting distribution ranges

In flowering plants, pollinators contribute to gene flow while they also respond to variation in plant traits together determined by genetic, epigenetic and environmental sources of variation. Consequently, a correlation between abundance and diversity of pollinators and the genetic and epigenetic characteristics of plant populations such as diversity or distinctiveness is to be expected, yet no study has explored such long-term dimensions of plant-pollinator interactions. Mediterranean narrow endemics often exhibit unexpectedly high levels of population (epi)genetic diversity. We hypothesize that pollinators may contribute to explain this pattern. Specifically, we expect a stronger association of pollinators with population (epi)genetic variability in narrow endemics than in widely distributed congeners. We studied five pairs of congeneric plant species, consisting of one narrow endemic with a restricted distribution and one widespread congener, found in the Sierra de Cazorla mountains (SE Spain). In up to three populations per species, a comprehensive characterisation of pollinators was carried out to estimate pollinator diversity and visitation rate. Genetic and epigenetic diversity and distinctiveness of each population was calculated using AFLP markers and methylation-sensitive AFLP markers (MSAP) respectively. The relationships with pollinator diversity and visitation rate were assessed. The diversity of pollinators did not vary according to the plant distribution range, whereas visitation rate was higher in widespread species. As predicted, only narrow endemics showed a significant association between pollinators and their population genetic and epigenetic characteristics. Specifically, higher pollinator diversity and visitation rates entailed higher population genetic diversity and lower epigenetic distinctiveness. This work shows the value of exploring the relationships between pollinator abundance and diversity and population (epi)genetics for understanding the evolution of plant rarity.

ecology↗

Comparative epigenetic and genetic spatial structure in Mediterranean mountain plants: a multispecific comparison

AO_SCPLOWBSTRACTC_SCPLOWEpigenetic information can be heritable, but also respond to key environmental variables in situ, endowing individuals with an additional capacity to adapt to environmental changes. Thus, it is likely that in sesile organisms such as plants, part of the spatial epigenetic variation found across individuals will reflect the environmental heterogeneity of populations. Analysing the departure of the spatial epigenetic structure from the baseline genetic variation can help in understanding the value of epigenetic regulation in species with different breath of optimal environmental conditions. We performed a multispecies study that considered seven pairs of congeneric plant species, each encompassing a narrow endemic with habitat specialization and a widespread species. In three populations per species we used AFLP and methylation-sensitive AFLP markers to characterise the spatial genetic and epigenetic structures. In contrast to widespread species, narrow endemics showed a significant and generalised lower epigenetic than genetic differentiation across species. Within most populations of narrow species, epigenetic variation was less spatially structured than the genetic variation. This pattern resulted from a lack of correlation between epigenetic and genetic information in populations of narrow endemics. We argue that the differences found between narrow endemics and widespread congeners reflect contrasting breaths of environmental requirements. We pose the hypothesis that in species with a narrow niche breath, epigenetic variation may be more similar across populations and among individuals within a population given the expected higher similarity in environmental requirements.

genetics↗

Body mass decline in a Mediterranean community of solitary bees supports the size shrinking effect of climatic warming

The long-known, widely documented inverse relationship between body size and environmental temperature ("temperature-size rule") has recently led to predictions of body size decline following current climatic warming ("size shrinking effect"). For keystone pollinators such as wild bees, body shrinking in response to warming can have pervasive effects on pollination processes, but there is still little evidence of the phenomenon because adequate tests require controlling for climate-linked confounding factors (e.g., urbanization, land use change). This paper tests the shrinking effect in a diverse community of solitary bees from well-preserved habitats in the core of a large nature reserve undergoing fast climatic warming but not disturbances or habitat changes. Long-term variation in mean body mass was evaluated using data from 1,186 individual bees (108 species, 25 genera, 6 families) sampled over 1990-2022. Climate of the region warmed at a fast rate during this period and changes in bee body mass verified expectations from the size shrinking effect. Mean individual body mass of the regional community of solitary bees declined significantly, shrinking being particularly intense for female individuals and large-bodied species. As a consequence, the pollination and mating systems of many bee-pollinated plants in the region are likely undergoing important alterations.

ecology↗

Seasonality of pollinators in Mediterranean montane habitats: cool-blooded bees for early-blooming plants

Understanding the factors that drive community-wide assembly of plant-pollinator systems along environmental gradients has considerable evolutionary, ecological and applied significance. Variation in thermal environments combined with intrinsic differences among pollinators in thermal biology (tolerance limits, thermal optima, thermoregulatory ability) have been proposed as drivers of community-wide pollinator gradients, but this suggestion remains largely speculative. We test the hypothesis that seasonality in bee pollinator composition in montane habitats of southeastern Spain, which largely reflects the prevalence during the early flowering season of mining bees (Andrena), is a consequence of the latters thermal biology. Quantitative information on seasonality of Andrena bees in the whole plant community (275 plant species) was combined with field and laboratory data on key aspects of the thermal biology of 30 species of Andrena (endothermic ability, warming constant, relationships of body temperature with ambient and operative temperatures). Andrena bees were a conspicuous, albeit strongly seasonal component of the pollinator assemblage of the regional plant community, visiting flowers of 153 different plant species (57% of total). Proportion of Andrena relative to all bees reached a maximum among plant species which flowered in late winter and early spring, and declined precipitously from May onwards. Andrena were recorded only during the cooler segment of the annual range of air temperatures experienced at flowers by the whole bee assemblage. These patterns can be explained by features of Andrenas thermal biology: null or negligible endothermy; ability to forage at much lower body temperature than endothermic bees (difference ~10{degrees}C); low upper tolerable limit of body temperature, beyond which thermal stress presumably precluded foraging at the warmest period of year; weak thermoregulatory capacity; and high warming constant enhancing ectothermic warming. Our results demonstrate the importance of lineage-specific pollinator traits as drivers of seasonality in community-wide pollinator composition; show that exploitation of cooler microclimates by bees does not require endothermy; falsify the frequent assumption that endothermy and thermoregulation apply to all bees; and suggest that medium- and large-sized ectothermic bees with low upper thermal limits and weak thermoregulatory ability can actually be more adversely affected by climate warming than large endothermic species.

ecology↗

Lifetime genealogical divergence within plants leads to epigenetic mosaicism in the long lived shrub Lavandula latifolia (Lamiaceae)

O_LIEpigenetic 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. C_LIO_LILeaf 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 epigenetic traits. C_LIO_LIPlants were epigenetically heterogeneous, as shown by differences among modules in global DNA methylation and variation in the methylation states of 6-8% of MS-AFLP markers. All epigenetic features exhibited significant genealogical signal within plants. Events of epigenetic divergence occurred throughout the lifespan of individuals and were subsequently propagated by branch divisions. C_LIO_LIInternal epigenetic diversification of L. latifolia individuals took place steadily during their development, a process which eventually led to persistent epigenetic mosaicism. C_LI

plant biology↗

Unclusterable, underdispersed arrangement of plant species in pollinator niche space

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 spaces were considered whose axes were defined by insect orders, families, genera and species; and (2) If all plant species form a single, indivisible cluster in pollinator niche space, Are they overdispersed or underdispersed relative to a random arrangement ? "Clusterability" tests failed to reject the null hypothesis that there was only one pollinator-defined plant species cluster in pollinator niche space, irrespective of spatial scale, pollinator niche space or pollinator importance measurement (proportions of pollinator individuals or flowers visited by each pollinator type). Observed means of pairwise interspecific dissimilarity in pollinator composition were smaller than randomly simulated values in the order-, family- and genus-defined pollinator niche spaces at both spatial scales, thus revealing significantly non-random, underdispersed arrangement of plant species within the single cluster existing in each of these pollinator niche spaces. In the undisturbed montane habitats studied, arrangement of insect-pollinated plant species in pollinator niche space did not support a major role for interspecific competition as a force shaping community-wide pollinator resource use by plants, but rather suggested a situation closer to the facilitation-dominated extreme in a hypothetical competition-facilitation gradient. Results also highlight the importance of investigations on complete or nearly complete insect-pollinated plant communities for addressing novel hypotheses on the ecology and evolution of plant-pollinator systems.

ecology↗

Gradual replacement of wild bees by honeybees in flowers of the Mediterranean Basin over the last 50 years

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 were gradually replaced by honeybees in flowers of wild and cultivated plants. Proportion of wild bees at flowers quadruplicated that of honeybees at the beginning of the period, the proportions of both groups becoming roughly similar fifty years later. The Mediterranean Basin is a world biodiversity hotspot for wild bees and wild bee-pollinated plants, and the ubiquitous rise of honeybees to dominance as pollinators could in the long run undermine the diversity of plants and wild bees in the region.

ecology↗