Global gradients in intraspecific variation in vegetative and floral traits are partially associated with climate and species richness

The study would not have been possible without the work of the TRY initiative on plant traits (http://www.try-db.org) and the BIEN database (http://bien.nceas.ucsb.edu/bien/). We thank all the BIEN and TRY contributors. Furthermore, we thank Benjamin Blonder, Melanie Harze, Ruben Milla, Clara Pladevall, Quentin Read, Marko Spasojevic, Alexia Totte, Evan R. Weiher, Ian Wright and Gerhard W. Zotz for providing additional information for their datasets. Additionally, we thank Franziska Schrodt and anonymous reviewers for constructive and thoughtful suggestions on earlier versions of this paper. Open Access funding was provided by Ulm University under the DEAL-agreement.Aim: Intraspecific trait variation (ITV) within natural plant communities can be large, influencing local ecological processes and dynamics. Here, we shed light on how ITV in vegetative and floral traits responds to large-scale abiotic and biotic gradients (i.e., climate and species richness). Specifically, we tested whether associations of ITV with temperature, precipitation and species richness were consistent with any of four hypotheses relating to stress tolerance and competition. Furthermore, we estimated the degree of correlation between ITV in vegetative and floral traits and how they vary along the gradients. Location: Global. Time period: 1975–2016. Major taxa studied: Herbaceous and woody plants. Methods: We compiled a dataset of 18,401 measurements of the absolute extent of ITV (measured as the coefficient of variation) in nine vegetative and seven floral traits from 2,822 herbaceous and woody species at 2,372 locations. Results: Large-scale associations between ITV and climate were trait specific and more prominent for vegetative traits, especially leaf morphology, than for floral traits. The ITV showed pronounced associations with climate, with lower ITV values in colder areas and higher values in drier areas. The associations of ITV with species richness were inconsistent across traits. Species-specific associations across gradients were often idiosyncratic, and covariation in ITV was weaker between vegetative and floral traits than within the two trait groups. Main conclusions: Our results show that, depending on the traits considered, ITV either increased or decreased with climate stress and species richness, suggesting that both factors can constrain or enhance ITV, which might foster plant-population persistence in stressful conditions. Given the species-specific responses and covariation in ITV, associations can be hard to predict for traits and species not yet studied. We conclude that consideration of ITV can improve our understanding of how plants cope with stressful conditions and environmental change across spatial and biological scales

Promoting individual and collective creativity in science students

Creativity is a scientific skill necessary to develop a successful research career. We expose the importance of a growth mindset, divergent, lateral, and associative thinking, serendipity, and being part of a nonhierarchical and diverse research team to improve both indi- vidual and collective creativity.Unidad de Excelencia Modeling Nature. Proyecto SOMM17/6109/UGR Consejería de Economía, Conocimiento, Empresas y Universidad, and European Regional Development Fund (ERDF

Genetic evidences of natural selection

La selección natural es un proceso biológico que constituye uno de los principales motores de cambio evolutivo, y el origen de las adaptaciones fenotípicas. La selección que actúa sobre una determinada población se puede detectar ecológicamente cuantificando a nivel intrapoblacional la relación existente entre el fenotipo de los individuos y su éxito reproductivo diferencial. Sin embargo, detectar la actuación de la selección pretérita es imposible usando una aproximación exclusivamente ecológica. Para afrontar este problema, durante los últimos 40 años se han desarrollado una serie de métodos moleculares que infieren la actuación de la selección. Un primer grupo de pruebas usa como modelo nulo la teoría cuasi neutral de la evolución molecular, y asumen que ha ocurrido selección cuando el resultado difiere significativamente de lo esperable según evolución neutral. Algunos tests, como por ejemplo la D de Tajima, la relación dN/dS , o las pruebas de MK o HKA, pueden determinar incluso el tipo de selección operante. Sin embargo, raramente pueden cuantificar la intensidad de selección. Este inconveniente es exitosamente superado por el análisis de la selección basada en Campos Aleatorios de Poisson (PRF). Un segundo grupo de pruebas usa como método para inferir la acción pretérita de la selección natural la comparación entre la cantidad de diferenciación genética en caracteres cuantitativos (QST) y la cantidad de diferenciación genética neutra (FST). Finalmente, unos de los más recientes y prometedores métodos que se están desarrollando para el estudio de la selección natural está relacionado con el avance tan espectacular que está registrando en la actualidad la genómica. Aún estamos lejos de saber la verdadera utilidad de esta herramienta para el estudio de la selección natural en poblaciones naturales, pero el pronóstico es bastante esperanzador.Natural selection is one of the main factors driving the evolutionary process and the only one leading to adaptations. Selection acting on a population can be detected by quantifying the relationship between individual phenotype and relative fitness. However, detecting the effect of past selection is not possible by only using an ecological approach. To cope with this issue, several molecular evolutionary genetics methods have been developed during the last 40 years. A first group of tests use the quasi neutral theory of molecular evolution as null model, and assume the occurrence of selection when the observed outcome significantly departs from the expectations under neutral evolution. Some tests, such as Tajima’s D, dN/dS ratio, MK test or HKA test, can determine the kind of selection acting on populations, but are unable to quantify its intensity. The analysis of selection based on Poisson Random Fields (PRF) overcomes this caveat. A second group of analyses infer selection by comparing the amount of genetic variation in quantitative traits (QST) against the amount of neutral genetic variation (FST). Lastly, one of the most recent and promising methods to explore natural selection is related with the superb advance in genomics. Although we are still far from fully appreciate the importance of this tool, we believe that the use of the genomics will produce a qualitative enhancement in our understanding of the importance of natural selection in the wild

Genomic Resources for Erysimum spp. (Brassicaceae): Transcriptome and Chloroplast Genomes

Funding was provided by the Spanish Ministry of Science and Competitiveness (CGL2016-79950-R; CGL2017-86626-C22-P), including FEDER funds. This research was also funded by the Consejeria de Economia, Conocimiento, Empresas y Universidad, and European Regional Development Fund (ERDF), ref. SOMM17/6109/UGR and A-RNM-505-UGR18. COM was supported by the Ministry of Economy and Competitiveness (BES-2014-069022).We are grateful to Modesto Berbel Cascales and José M. Gómez for their help in sampling and DNA/RNA extractions.Erysimum (Brassicaceae) is a genus of more than 200 species (Al-Shehbaz, 2012). It is widely distributed in the Northern Hemisphere and has been the focus of active research in ecology, evolution, and genetics (Gómez and Perfectti, 2010; Gómez, 2012; Valverde et al., 2016). Despite long-standing interest in Erysimum, its taxonomy has yet to be properly established, partly due to a complex and reticulated evolutionary history that renders phylogenetic reconstructions highly challenging (Ancev, 2006; Marhold and Lihová, 2006; Abdelaziz et al., 2014; Gomez et al., 2014; Moazzeni et al., 2014; Züst et al., 2020). The Baetic Mountains (South-Eastern Iberia) are among the most critical glacial refugia in Europe. The waxing and waning of plant populations following climatic fluctuations have likely complicated the distribution and genetic variation of extant diversity in this region. Isolation and posterior secondary contact between taxa may have favored hybridization and introgression (Médail and Diadema, 2009). The Erysimum species that inhabit these mountains have been a particularly fruitful system for plant evolutionary ecology [e.g., Gómez et al., 2006, 2008; Gómez and Perfectti, 2010; Gómez, 2012; Valverde et al., 2016]. However, the relationships among these species remain unresolved, hampering comparative and evolutionary studies. Genome duplications, incomplete lineage sorting, and hybridization have compromised the phylogenetic reconstructions within Erysimum (Marhold and Lihová, 2006; Osuna-Mascaró, 2020). Additionally, clarifying this group's complex evolution requires extensive genomic resources, which are currently being produced but are mostly lacking. The fast development of high-throughput sequencing technologies has led to a rapid increase in genomic and transcriptomic for many plant species (Dong et al., 2004; Duvick et al., 2007; Sundell et al., 2015; Boyles et al., 2019). However, obtaining complete genome sequencing remains a challenge with large, repetitive-DNA enriched genomes. Transcriptome sequencing is comparatively more accessible, providing a relatively cheap and fast method to obtain large amounts of functional genomic data (Timme et al., 2012; Yang and Smith, 2013; Wickett et al., 2014; Léveillé-Bourret et al., 2017). Accordingly, global initiatives such as the 1,000 plants (1KP) project have generated transcriptomic resources for over 1,000 plant species (Matasci et al., 2014; Leebens-Mack et al., 2019). In addition, the use of RNA-Seq could be useful in obtaining complete chloroplast genomes in a reliable and accessible way, making possible the use of complete molecules in phylogenomic analyses (Smith, 2013; Osuna-Mascaró et al., 2018; Morales-Briones et al., 2021). Here, we report the annotation of 18 floral transcriptomes assembled de novo from total RNA-Seq libraries and nine chloroplast genomes from seven Erysimum species inhabiting the Baetic Mountains. The chloroplast genomes were assembled from total RNA-Seq data following a previously-validated reference assemble approach (Osuna-Mascaró et al., 2018). The data presented here represent reliable genomic resources for transcriptomic, proteomic, and phylotranscriptomic studies. These data contribute to the ecological and genetic resources available for Brassicaceae in general and the genus Erysimum in particular, being the only genomic resources for these species coming from flower buds.Spanish Ministry of Science and Competitiveness CGL2016-79950-R CGL2017-86626-C22-PEuropean CommissionConsejeria de Economia, Conocimiento, Empresas y UniversidadEuropean Commission SOMM17/6109/UGR A-RNM-505-UGR18Ministry of Economy and Competitiveness BES-2014-06902

The role of phenotypic plasticity in shaping ecological networks

Plasticity-mediated changes in interaction dynamics and structure may scale up and affect the ecological network in which the plastic species are embedded. Despite their potential relevance for understanding the effects of plasticity on ecological communities, these effects have seldom been analysed. We argue here that, by boosting the magnitude of intra-individual phenotypic variation, plasticity may have three possible direct effects on the interactions that the plastic species maintains with other species in the community: may expand the interaction niche, may cause a shift from one interaction niche to another or may even cause the colonization of a new niche. The combined action of these three factors can scale to the community level and eventually expresses itself as a modification in the topology and functionality of the entire ecological network. We propose that this causal pathway can be more widespread than previously thought and may explain how interaction niches evolve quickly in response to rapid changes in environmental conditions. The implication of this idea is not solely eco-evolutionary but may also help to understand how ecological interactions rewire and evolve in response to global changeJunta de Andalucía, Grant/Award Number: P18-FR- 3641Spanish Ministry of Science, Innovation and UniversitiesGrant/Award Number: PID2020-116222GB- 100PID2021-126456N

Phenotypic plasticity guides Moricandia arvensis divergence and convergence across the Brassicaceae floral morphospace

Authors thank Raquel Sánchez, Angel Caravantes, Isabel Sánchez Almazo, María José Jorquera, and Iván Rodríguez Arós for helping us during several phases of the study. We also thank all contributors to the pollinator database (Table  S1 ) for kindly sending us unpublished information on Brassicaceae floral visitors. This research is supported by grants from the Spanish Ministry of Science, Innovation and Universities (CGL2015‐63827‐P, CGL2017‐86626‐C2‐1‐P, CGL2017‐86626‐C2‐2‐P, UNGR15‐CE‐3315), Junta de Andalucía (P18‐FR‐3641, IE19_238 EEZA CSIC), LIFE18 GIE/IT/000755, and Xunta de Galicia (CITACA), including EU FEDER funds. This is a contribution to the Research Unit Modeling Nature, funded by the Consejería de Economía, Conocimiento, Empresas y Universidad, and European Regional Development Fund (ERDF), reference SOMM17/6109/UGR.Many flowers exhibit phenotypic plasticity. By inducing the production of several phenotypes, plasticity may favour the rapid exploration of different regions of the floral morphospace. We investigated how plasticity drives Moricandia arvensis, a species displaying within-individual floral polyphenism, across the floral morphospace of the entire Brassicaceae family. We compiled the multidimensional floral phenotype, the phylogenetic relationships, and the pollination niche of over 3000 species to construct a family-wide floral morphospace. We assessed the disparity between the two M. arvensis floral morphs (as the distance between the phenotypic spaces occupied by each morph) and compared it with the family-wide disparity. We measured floral divergence by comparing disparity with the most common ancestor, and estimated the convergence of each floral morph with other species belonging to the same pollination niches. Moricandia arvensis exhibits a plasticity-mediated floral disparity greater than that found between species, genera and tribes. The novel phenotype of M. arvensis moves outside the region occupied by its ancestors and relatives, crosses into a new region where it encounters a different pollination niche, and converges with distant Brassicaceae lineages. Our study suggests that phenotypic plasticity favours floral divergence and rapid appearance of convergent flowers, a process which facilitates the evolution of generalist pollination systems.CITACAConsejería de Economía, Conocimiento, Empresas y UniversidadMinisterio de Ciencia, Innovación y Universidades UNGR15‐CE‐3315European Regional Development FundXunta de GaliciaJunta de Andalucía IE19_238 EEZA CSIC, LIFE18 GIE/IT/00075

Spread of a New Parasitic B Chromosome Variant Is Facilitated by High Gene Flow

The B24 chromosome variant emerged several decades ago in a Spanish population of the grasshopper Eyprepocnemis plorans and is currently reaching adjacent populations. Here we report, for the first time, how a parasitic B chromosome (a strictly vertically transmitted parasite) expands its geographical range aided by high gene flow in the host species. For six years we analyzed B frequency in several populations to the east and west of the original population and found extensive spatial variation, but only a slight temporal trend. The highest B24 frequency was found in its original population (Torrox) and it decreased closer to both the eastern and the western populations. The analysis of Inter Simple Sequence Repeat (ISSR) markers showed the existence of a low but significant degree of population subdivision, as well as significant isolation by distance (IBD). Pairwise Nem estimates suggested the existence of high gene flow between the four populations located in the Torrox area, with higher values towards the east. No significant barriers to gene flow were found among these four populations, and we conclude that high gene flow is facilitating B24 diffusion both eastward and westward, with minor role for B24 drive due to the arrival of drive suppressor genes which are also frequent in the donor population.This study was supported by a grant from the Spanish Ministerio de Ciencia e Innovación (CGL2009-11917), and was partially performed by FEDER ("Fondo Europeo de Desarrollo" - European Regional Development Fund - ERDF) funds. MIMP was supported by a fellowship (FPU) from the Spanish Ministerio de Ciencia e Innovación

Geographical Barriers Impeded the Spread of a Parasitic Chromosome

Parasitic supernumerary (B) chromosomes show high capability to spread across populations. But the existence of abrupt discontinuities in their distribution demands an explanation. The grasshopper Eyprepocnemis plorans plorans harbour supernumerary chromosomes in all natural populations hitherto analyzed from the Circum-Mediterranean region, with the single exception of the headwaters of the Iberian Segura River and several of its tributaries. To ascertain the causes of this distribution pattern, we analyze here the genetic structure of five natural populations collected in this zone (two +B and three -B), by means of ISSR markers. We found significant population structure, with two kinds of populations coinciding with +B and -B ones, separated by strong barriers to gene flow. This gives strong support to the hypothesis that the non-B populations precede B origin, and that B-carrying individuals from coastal zones have been able to colonize upstream areas, until geographical barriers (usually narrow canyons and arid areas surrounding them) impeded their advance.The authors received no specific funding for this work. Project applications have been rejected in 2012, 2013 and 2014

Interpopulation spread of a parasitic B chromosome is unlikely through males in the grasshopper Eyprepocnemis plorans

This study was supported by grants from the Spanish Secretaría de Estado de Investigación, Desarrollo e Innovación (CGL2015-70750-P), and was partially performed by FEDER funds.The near-neutral model of B chromosome evolution predicts that population invasion is quite fast. To test this prediction, in 1994, we introduced males of the grasshopper Eyprepocnemis plorans from a B-carrying population into a B-lacking population and monitored the evolution of B-chromosome frequency up to 2013. We observed fluctuating very low B frequency across years but, remarkably, the B chromosome introduced (the B2 variant) was found up to 1996 only, whereas the B1 variant was present from 1996 onwards, presumably introduced by fishermen using E. plorans males as bait. Effective introgression of genetic material from the donor population was evidenced by the presence of a satellite DNA on autosome 9 (up to 1999) and the presence of one individual in 2006 showing an ISSR marker profile being highly similar to that found in the donor population. This indicated that the males introduced by us effectively mated with resident females, but donor genes rapidly decreased in frequency after this non-recurrent migration event. Taken together, our results indicated: (i) that the non-recurrent migration event had a slight, transient genetic effect on the recipient population, which was diluted in only a few generations; and (ii) that even with recurrent migration (forced by fishermen) the B chromosome failed to increase in frequency. Bearing in mind that B chromosomes in this species drive through females only, we hypothesize that B chromosomes most likely failed invasion in both migration events because the migrating sex shows no B-drive.Spanish Secretaría de Estado de Investigación, Desarrollo e Innovación (CGL2015-70750-P)FEDER fund

Within-individual phenotypic plasticity in flowers fosters pollination niche shift

Phenotypic plasticity, the ability of a genotype of producing different phenotypes when exposed to different environments, may impact ecological interactions. We study here how within-individual plasticity in Moricandia arvensis flowers modifies its pollination niche. During spring, this plant produces large, cross-shaped, UV-reflecting lilac flowers attracting mostly long-tongued large bees. However, unlike most co-occurring species, M. arvensis keeps flowering during the hot, dry summer due to its plasticity in key vegetative traits. Changes in temperature and photoperiod in summer trigger changes in gene expression and the production of small, rounded, UV-absorbing white flowers that attract a different assemblage of generalist pollinators. This shift in pollination niche potentially allows successful reproduction in harsh conditions, facilitating M. arvensis to face anthropogenic perturbations and climate change.Ministerio de Economía y Competitividad | Ref. CGL2015-71634-PMinisterio de Economía y Competitividad | Ref. CGL2015-63827-PAgencia Estatal de Investigación | Ref. CGL2017-86626-C2-1-PAgencia Estatal de Investigación | Ref. CGL2017-86626-C2-2-PMinisterio de Economía y Competitividadd | Ref. UNGR15-CE-3315Ministerio de Economía y Competitividad | Ref. RYC-2012-12277Ministerio de Economía y Competitividad | Ref. UNGR15-CE-3315Junta de Andalucía | Ref. P18-FR-3641Fundación BBVA | Ref. PR17_ECO_0021Xunta de Galici