Ploidy effects on the relationship between floral phenotype, reproductive investment, and fitness in an autogamous species complex.

PREMISE: The relationships between reproductive investment, phenotype, and fitness have been broadly studied in cross-pollinated plants in contrast to selfing species, which are considered less interesting in this area because they are supposed to be a dead end in any evolutionary pathway. Still, selfing plants are unique systems to study these questions since the position of reproductive structures and traits related to flower size play an important role in female and male pollination success. METHODS: Erysimum incanum s.l. is a selfing species complex that has three levels of ploidy (diploids, tetraploids, and hexaploids) and traits that are typically associated with the selfing syndrome. Here, we used 1609 plants belonging to these three ploidies to characterize the floral phenotype and spatial configuration of reproductive structures, reproductive investment (pollen and ovule production), and plant fitness. Then, we used structural equation modelling to analyze the relationship between all these variables across ploidy levels. RESULTS: An increase in ploidy level leads to bigger flowers with anthers exserted farther and more pollen and ovules. In addition, hexaploid plants had higher absolute values for herkogamy, which is positively correlated with fitness. Ovule production significantly mediated the natural selection acting on different phenotypic traits and pollen production, a pattern that is maintained across ploidies. CONCLUSIONS: The changes in floral phenotypes, reproductive investment, and fitness with ploidy level suggest that genome duplication can be a driver for transitions in reproductive strategy by modifying the investment in pollen and ovules and linking them with plant phenotype and fitness.

Direct evidence supporting Darwin's hypothesis of cross-pollination promoted by sex organ reciprocity.

The floral phenotype plays a main role in the attraction and fit of pollinators. Both perianth traits and the positioning of sex organs can be subjected to natural selection and determine nonrandom mating patterns in populations. In stylar-polymorphic species, the Darwinian hypothesis predicts increased mating success between individuals with sex organs at equivalent heights (i.e. with higher reciprocity). We used paternity analyses in experimental populations of a stylar-dimorphic species. By comparing the observed mating patterns with those expected under random mating, we tested the effects of sex organ reciprocity and perianth traits on mating success. We also analysed phenotypic selection on perianth traits through female and male functions. The (dis)similarity of parental perianth traits had no direct effects on the mating patterns. Sex organ reciprocity had a positive effect on mating success. Narrow floral tubes increased this effect in upper sex organs. Perianth traits showed little signs of phenotypic selection. Female and absolute fitness measures resulted in different patterns of phenotypic selection. We provide precise empirical evidence of the Darwinian hypothesis about the functioning of stylar polymorphisms, demonstrating that mating patterns are determined by sex organ reciprocity and only those perianth traits which are critical to pollinator fit.

Asymmetric Reproductive Barriers and Gene Flow Promote the Rise of a Stable Hybrid Zone in the Mediterranean High Mountain.

Hybrid zones have the potential to shed light on evolutionary processes driving adaptation and speciation. Secondary contact hybrid zones are particularly powerful natural systems for studying the interaction between divergent genomes to understand the mode and rate at which reproductive isolation accumulates during speciation. We have studied a total of 720 plants belonging to five populations from two Erysimum (Brassicaceae) species presenting a contact zone in the Sierra Nevada mountains (SE Spain). The plants were phenotyped in 2007 and 2017, and most of them were genotyped the first year using 10 microsatellite markers. Plants coming from natural populations were grown in a common garden to evaluate the reproductive barriers between both species by means of controlled crosses. All the plants used for the field and greenhouse study were characterized by measuring traits related to plant size and flower size. We estimated the genetic molecular variances, the genetic differentiation, and the genetic structure by means of the F-statistic and Bayesian inference. We also estimated the amount of recent gene flow between populations. We found a narrow unimodal hybrid zone where the hybrid genotypes appear to have been maintained by significant levels of a unidirectional gene flow coming from parental populations and from weak reproductive isolation between them. Hybrid plants exhibited intermediate or vigorous phenotypes depending on the analyzed trait. The phenotypic differences between the hybrid and the parental plants were highly coherent between the field and controlled cross experiments and through time. The highly coherent results obtained by combining field, experimental, and genetic data demonstrate the existence of a stable and narrow unimodal hybrid zone between Erysimum mediohispanicum and Erysimum nevadense at the high elevation of the Sierra Nevada mountains.

Temporal migration rates affect the genetic structure of populations in the biennial Erysimum mediohispanicum with reproductive asynchrony.

Migration is a process with important implications for the genetic structure of populations. However, there is an aspect of migration seldom investigated in plants: migration between temporally isolated groups of individuals within the same geographic population. The genetic implications of temporal migration can be particularly relevant for semelparous organisms, which are those that reproduce only once in a lifetime after a certain period of growth. In this case, reproductive asynchrony in individuals of the same population generates demes of individuals differing in their developmental stage (non-reproductive and reproductive). These demes are connected by temporal migrants, that is, individuals that become annually asynchronous with respect to the rest of individuals of their same deme. Here, we investigated the extent of temporal migration and its effects on temporal genetic structure in the biennial plant Erysimum mediohispanicum. To this end, we conducted two independent complementary approaches. First, we empirically estimated temporal migration rates and temporal genetic structure in four populations of E. mediohispanicum during three consecutive years using nuclear microsatellites markers. Second, we developed a demographic genetic simulation model to assess genetic structure for different migration scenarios differing in temporal migration rates and their occurrence probabilities. We hypothesized that genetic structure decreased with increasing temporal migration rates due to the homogenizing effect of migration. Empirical and modelling results were consistent and indicated a U-shape relationship between genetic structure and temporal migration rates. Overall, they indicated the existence of temporal genetic structure and that such genetic structure indeed decreased with increasing temporal migration rates. However, genetic structure increased again at high temporal migration rates. The results shed light into the effects of reproductive asynchrony on important population genetic parameters. Our study contributes to unravel the complexity of some processes that may account for genetic diversity and genetic structure of natural populations.

The progress of interdisciplinarity in invasion science.

Interdisciplinarity is needed to gain knowledge of the ecology of invasive species and invaded ecosystems, and of the human dimensions of biological invasions. We combine a quantitative literature review with a qualitative historical narrative to document the progress of interdisciplinarity in invasion science since 1950. Our review shows that 92.4% of interdisciplinary publications (out of 9192) focus on ecological questions, 4.4% on social ones, and 3.2% on social-ecological ones. The emergence of invasion science out of ecology might explain why interdisciplinarity has remained mostly within the natural sciences. Nevertheless, invasion science is attracting social-ecological collaborations to understand ecological challenges, and to develop novel approaches to address new ideas, concepts, and invasion-related questions between scholars and stakeholders. We discuss ways to reframe invasion science as a field centred on interlinked social-ecological dynamics to bring science, governance and society together in a common effort to deal with invasions.

PHENIX: An R package to estimate a size-controlled phenotypic integration index.

PREMISE OF THE STUDY: Organisms usually show intercorrelations between all or some of their components leading to phenotypic integration, which may have deep consequences on the evolution of phenotypes. One of the main difficulties with phenotypic integration studies is how to correct the integration measures for size. This has been considered a challenging task. In this paper, we introduce an R package (PHENIX: PHENotypic Integration indeX), in which we provide functions to estimate a size-controlled phenotypic integration index, a bootstrapping method to calculate confidence intervals, and a randomization method to simulate null distributions and test the statistical significance of the integration. METHODS AND RESULTS: PHENIX is an open source package written in R. As usual for R packages, the manual and sample data are available at: http://cran.r-project.org/web/packages/PHENIX/index.html. Functions included in this package easily estimate phenotypic integration by controlling a third variable (e.g., the size of the studied organ). CONCLUSIONS: PHENIX helps to estimate and test the statistical significance of the magnitude of integration using one of the most-used methodological approaches, while taking size into account.

Characterization of microsatellite loci in Erysimum mediohispanicum (Brassicaceae) and cross-amplification in related species.

PREMISE OF THE STUDY: We have developed and optimized microsatellite loci from a genomic library of Erysimum mediohispanicum. Microsatellites were also tested for cross-amplification in 31 other Erysimum species. METHODS AND RESULTS: A total of 10 microsatellite loci were successfully amplified. They were polymorphic for 81 E. mediohispanicum individuals from two locations in Sierra Nevada (southeastern Spain), which showed similar patterns of genetic diversity. On average, microsatellites had 8.6 alleles per locus and an expected heterozygosity of 0.69. Only one locus significantly departed from Hardy-Weinberg equilibrium in both locations. Most of the markers successfully amplified in other Erysimum species. CONCLUSIONS: The genetic attributes of microsatellite loci will allow their application to population genetic studies in Erysimum, such as genetic differentiation and structure, gene flow, pollinator-mediated speciation, and hybridization studies.

Using complementary techniques to distinguish cryptic species: a new Erysimum (Brassicaceae) species from North Africa.

PREMISE OF THE STUDY: Cryptic species are superficially morphologically indistinguishable and therefore erroneously classified under one single name. The identification and delimitation of these species is usually a difficult task. The main aim of this study is to provide an inclusive methodology that combines standard and new tools to allow accurate identification of cryptic species. We used Erysimum nervosum s.l. as a model system. METHODS: Four populations belonging to E. nervosum s.l. were sampled at their two distribution ranges in Morocco (the Atlas Mountains and the Rif Mountains). Fifteen individuals per population were collected to assess standard taxonomic traits. Additionally, corolla color and shape were quantified in 30 individuals per population using spectrophotometry and geometric morphometrics, respectively. Finally, we collected tissue samples from each population per species to study the phylogenetic relationships among them. KEY RESULTS: Using the standard taxonomic traits, we could not distinguish the four populations. Nonetheless, there were differences in corolla color and shape between plants from the two mountain ranges. The population differentiation based on quantitative morphological differences were confirmed and supported by the phylogenetic relationships obtained for these populations and the rest of the Moroccan Erysimum species. CONCLUSIONS: The joint use of the results obtained from standard taxonomic traits, quantitative analyses of plant phenotype, and molecular data suggests the occurrence of two species within E. nervosum s.l. in Morocco, one located in the Atlas Mountains (E. nervosum s.s.) and the other in the Rif Mountains (E. riphaeanum sp. nov.). Consequently, we suggest that combining quantitative and molecular approaches with standard taxonomy greatly benefits the identification of cryptic species.

A single, recent origin of the accessory B chromosome of the grasshopper Eyprepocnemis plorans.

B chromosomes are dispensable chromosomes found in >2000 eukaryotic species, usually behaving as genomic parasites. Most B chromosomes seem to be made up of the same kind of DNA sequences present in the A chromosomes. This sequence similarity makes it difficult to obtain specific molecular probes that may permit B-presence diagnosis without cytogenetic analysis. We have developed a sequence-characterized amplified region (SCAR) marker for B chromosomes in the grasshopper Eyprepocnemis plorans, which specifically amplifies a 1510-bp DNA fragment exclusively in B-carrying individuals. Fluorescent in situ hybridization and fiber FISH analyses showed that this marker is a tandemly repeated DNA sequence closely intermingled with 45S rDNA. PCR reactions showed the presence of SCAR-like sequences in the A chromosomes, but in two separate fragments, supporting the intraspecific origin of B chromosomes in this species. SCAR marker DNA sequence showed to be identical in B chromosome variants from several localities from Spain and Morocco, and it was very similar to those found in B chromosome variants from Greece and Armenia. This strongly suggests that this sequence was already present in the ancestral B chromosome of this species. In addition, the scarce sequence variation observed among several B variants from very distant populations suggests either a functional constraint or, more likely, a recent and unique origin for B chromosomes in this species.