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Aim: Understanding the historical and contemporaneous drivers of invasion success in island systems can decisively contribute to identifying sources and pathways that are more likely to give rise to new invaders. Based on a floristic-driven approach, we aimed at determining the origins of the invasive alien flora of the Canary Islands and shedding light in the mechanisms shaping their distribution within the archipelago., This research was supported by the Fundación BBVA project (INVASION - PR19_ECO_0046) and the Spanish Ministry of Science and Innovation (MICINN) project (ASTERALIEN - PID2019-110538GA-I00). J.M.-L. was funded by the INVASION Fundación BBVA project and by a Juan de la Cierva-Formación Fellowship (MICIIN; reference FJC2020-046353-I). Y.A. by the ASTERALIEN MICINN project. J.P. was funded by the MICINN through the Ramón y Cajal Program (RYC-2016-20506). L.S.J.G. was funded by the Academia Canaria de Investigación Gobierno de Canarias FPI 2021 Fellowship (TESIS2021010101). The authors also thank Dr. D. Chapman and two anonymous reviewers for their constructive comments and suggestions. No permits were needed to perform this study.

A recurring feature of oceanic archipelagos is the presence of adaptive radiations that generate endemic, species-rich clades that can offer outstanding insight into the links between ecology and evolution. Recent developments in evolutionary genomics have contributed towards solving long-standing questions at this interface. Using a comprehensive literature search, we identify studies spanning 19 oceanic archipelagos and 110 putative adaptive radiations, but find that most of these radiations have not yet been investigated from an evolutionary genomics perspective. Our review reveals different gaps in knowledge related to the lack of implementation of genomic approaches, as well as undersampled taxonomic and geographic areas. Filling those gaps with the required data will help to deepen our understanding of adaptation, speciation, and other evolutionary processes., D.D.C. was supported by a Humboldt postdoctoral fellowship. J.C. was supported by a Research Council of Norway travel grant (project 287327). J.C. thanks Andy Foote for early input and ideas on what a review encompasses. The authors are extremely grateful to four anonymous reviewers and the editor whose thoughtful criticism led to a significant improvement of this manuscript. This publication is contribution number 2488 of the Charles Darwin Foundation for the Galapagos Islands.

Neutral theory proposes that dispersal stochasticity is one of the main drivers of local diversity. Haplotypes-level genetic variation can now be efficiently sampled from across whole communities, thus making it possible to test neutral predictions from the genetic to species-level diversity, and higher. However, empirical data is still limited, with the few studies to date coming from temperate latitudes. Here, we focus on a tropical mountain within the Transmexican Volcanic Belt to evaluate spatially fine-scale patterns of arthropod community assembly to understand the role of dispersal limitation and landscape features as drivers of diversity. We sampled whole-communities of arthropods for eight orders at a spatial scale ranging from 50 m to 19 km, using whole community metabarcoding. We explored multiple hierarchical levels, from individual haplotypes to lineages at 0.5, 1.5, 3, 5, 7.5% similarity thresholds, to evaluate patterns of richness, turnover, and distance decay of similarity with isolation-by-distance and isolation-by resistance (costs to dispersal given by landscape features) approaches. Our results showed that distance and altitude influence distance decay of similarity at all hierarchical levels. This holds for arthropod groups of contrasting dispersal abilities, but with different strength depending on the spatial scale. Our results support a model where local-scale differentiation mediated by dispersal constraints, combined with long-term persistence of lineages, is an important driver of diversity within tropical sky islands., Nancy Gálvez-Reyes is a doctoral student from Programa de Doctorado en Ciencias Biomédicas, Universidad Nacional Autónoma de México (UNAM) and was supported by CONACYT (scholarship no. 401781). CONACYT also provided financial support with the project 178245 granted to D. Piñero. B. Emerson was supported by the Spanish Agencia Estatal de Investigación (CGL2017-85718-P), co-financed by FEDER. P. Arribas was supported by a postdoctoral grant by the Spanish Ministry of Economy and Competitiveness (MINECO, Spain) within the Juan de la Cierva Formación Program. C. Andújar was supported by the Spanish Ministry of Economy and Competitiveness (MINECO, Spain) (CGL2015-74178-JIN MINECO/FEDER, UE).

In non-fire prone ecosystems, like some subtropical humid forests, fire produces habitat destruction and intensifies land degradation by inducing changes in native species composition, soil properties and erosive processes. Bryophytes are key components of the Macaronesian laurel forests playing an important role in regulating water cycling and microclimate. Ecological and taxonomical bryophytes groups have distinct ecological and physiological requirements and may respond in a different way to the same fire events. Therefore, analysing post fire recovery of bryophyte communities represents a key step towards a better understanding of forest fire drivers and post fire management. We investigated how species richness and composition of different ecological and taxonomical bryophyte groups varied in 1158 samples within a fire chronosequence from 5 to 57 years in the best-preserved laurel forest from Canary Islands (Garajonay National Park) analysing communities in terms of differences with comparable surrounding old growth unburnt stands. Epiphyte, terricolous and saxicolous bryophytes were sampled at each plot and the influence of the time since fire was analyzed together with environmental variables (temperature, precipitation, mist precipitation and elevation) and forest structure variables. Our results indicate that there is no general pattern of post fire recolonization, as recolonization varies depending on the ecological and phylogenetic groups considered. Climate and forest structure play an important role in post-fire recolonization, such that time since fire is not the most important variable influencing richness and composition. The results increase the understanding of the processes that shape compositional patterns in groups with high dispersal capacities and high microclimate dependence, such as mosses and liverworts., We are especially grateful to Julio Leal Pérez, for his help and assistance during all the fieldwork. We thank to Ángel Fernández López, from Garajonay National Park for their help in the fieldwork and for providing authorization and support people: especially Israel Rodríguez Reverón, Genaro Barrera and Luis Gómez. Many thanks also to Jonay Cubas, Ángel Mallorquín and Jairo Patiño, for providing help during the field work. Víctor Bello Rodríguez provided cartography data of La Gomera and relevant information on environmental variables. We also thank the funding of this research project titled “Cuantificando la respuesta a los incendios forestales de las comunidades de plantas y artrópodos de los bosques de laurisilva del Parque Nacional de Garajonay” by the Spanish Ministerio de Agricultura, Alimentación y Pesca and Organismo Autónomo de Parques Nacionales, Grant no. S20141203_002597. First author thanks the financial support from Agencia Canaria de Investigación, Innovación y Sociedad de la Información de la Consejería de Economía, Conocimiento y Empleo cofunded in a 85 % by the European Social Fund (ESF) to develop his PhD degree.

Current understanding of ecological and evolutionary processes underlying island biodiversity is heavily shaped by empirical data from plants and birds, although arthropods comprise the overwhelming majority of known animal species, and as such can provide key insights into processes governing biodiversity. Novel high throughput sequencing (HTS) approaches are now emerging as powerful tools to overcome limitations in the availability of arthropod biodiversity data, and hence provide insights into these processes. Here, we explored how these tools might be most effectively exploited for comprehensive and comparable inventory and monitoring of insular arthropod biodiversity. We first reviewed the strengths, limitations and potential synergies among existing approaches of high throughput barcode sequencing. We considered how this could be complemented with deep learning approaches applied to image analysis to study arthropod biodiversity. We then explored how these approaches could be implemented within the framework of an island Genomic Observatories Network (iGON) for the advancement of fundamental and applied understanding of island biodiversity. To this end, we identified seven island biology themes at the interface of ecology, evolution and conservation biology, within which collective and harmonized efforts in HTS arthropod inventory could yield significant advances in island biodiversity research., The iGON Working Group was organized by the iBioGen project, which has received funding from the European Union's Horizon 2020 research and innovation programme under grant agreement no. 810729.

Monitoring biodiversity is essential to protect, preserve and restore ecosystems, particularly in the context of current challenges such as climate change, habitat destruction and globalization (Baird & Hajibabaei, 2012). Biomonitoring is needed for developing biotic indices for assessing ecological status, measuring impacts of anthropogenic activities in natural ecosystems, evaluating biodiversity loss, surveying nonindigenous species, conservation, and identifying cryptic species (Balvanera et al., 2006; Fišer et al., 2018). Thus, spatially and temporally structured biomonitoring activities provide a powerful tool for the implementation of regional, national and international regulations, directives and policies for nature conservation. However substantial impediments exist including access to remote locations, limited specialist taxonomic knowledge, cost, slow pace of human-driven data analyses, and typically low sensitivity for detection of rare and elusive species (Zinger et al., 2020). These drawbacks are often translated into expensive monitoring activities with limited spatial, temporal and taxonomic coverage. In this context, new approaches for biomonitoring are being explored, among which advanced DNA-based technologies are emerging (Kissling et al., 2018). The field of biodiversity monitoring through the analysis of the pool of DNA isolated from environmental samples, referred to as environmental DNA or eDNA (Pawlowski et al., 2020; Taberlet et al., 2012), is rapidly growing. This growth is being driven through improved approaches for sampling, data generation and analyses, and with recent advances on how eDNA should be interpreted for biodiversity assessments (Bohmann et al., 2014). The success of eDNA-based biomonitoring is reflected in exponential growth of publications within this area and increasing submissions to Molecular Ecology Resources in particular (Figure 1). Molecular Ecology Resources aims to publish high quality eDNA studies that serve as broad resources, including innovative methodologies for DNA sampling, enhanced laboratory protocols for data generation, or new computer programs and statistical advances for data analyses. Thus, the aim of this editorial is to contribute to producing good quality DNA data-derived essential biodiversity variables (EBVs) (Kissling et al., 2018) by providing guidance to the community submitting articles on the subject. For that purpose, we have summarized best practices established in published literature related to the different phases involved in the process, namely sampling, laboratory work, bioinformatic analyses and data interpretation (Figure 2).

Most of our understanding of island diversity comes from the study of aboveground systems, while the patterns and processes of diversification and community assembly for belowground biotas remain poorly understood. Here, we take advantage of a relatively young and dynamic oceanic island to advance our understanding of ecoevolutionary processes driving community assembly within soil mesofauna. Using whole organism community DNA (wocDNA) metabarcoding and the recently developed metaMATE pipeline, we have generated spatially explicit and reliable haplotype-level DNA sequence data for soil mesofaunal assemblages sampled across the four main habitats within the island of Tenerife. Community ecological and metaphylogeographic analyses have been performed at multiple levels of genetic similarity, from haplotypes to species and supraspecific groupings. Broadly consistent patterns of local-scale species richness across different insular habitats have been found, whereas local insular richness is lower than in continental settings. Our results reveal an important role for niche conservatism as a driver of insular community assembly of soil mesofauna, with only limited evidence for habitat shifts promoting diversification. Furthermore, support is found for a fundamental role of habitat in the assembly of soil mesofauna, where habitat specialism is mainly due to colonization and the establishment of preadapted species. Hierarchical patterns of distance decay at the community level and metaphylogeographical analyses support a pattern of geographic structuring over limited spatial scales, from the level of haplotypes through to species and lineages, as expected for taxa with strong dispersal limitations. Our results demonstrate the potential for wocDNA metabarcoding to advance our understanding of biodiversity., This work was supported by projects CGL2015-74178-JIN (AEI, Spain/FEDER, EU) and SOILBIODIV (H2020-MSCA-IF-2015; award no.: 705639) awarded to CA, CGL2017-85718-P and PID2020-116788GB-I00 (AEI, Spain/FEDER, EU) and 2017RCE03 (Fundación CajaCanarias/Obra Social “La Caixa”) awarded to BCE. PA was additionally funded through a Junior Leader Fellowship (LCF/BQ/PR21/11840006) by “la Caixa” Foundation (ID 100010434) and the European Union's Horizon 2020 research and innovation programme under the Marie Skłodowska-Curie grant agreement No847648. We extend our gratitude to the regional governments of Andalucía, Canarias, and the local council (Cabildo) of Tenerife for facilitating collecting of samples, and to Jesús Arribas for assistance with field sampling and drawings of habitats.

In Focus: Scalercio, S., Cini, A., Menchetti, M., Vodă, R., Bonelli, S., Bordoni, A., … Dapporto, L. (2020). How long is 3 km for a butterfly? Ecological constraints and functional traits explain high mitochondrial genetic diversity between Sicily and the Italian Peninsula. Journal of Animal Ecology. https://doi.org/10.1111/1365‐2656.13196. Biotic and abiotic factors can shape geographical patterns of genetic variation within species, but few studies have addressed how this might generate common patterns at the level of communities of species. Scalercio et al. (2020) have combined mtDNA sequence data and life‐history traits, to reveal a repeated pattern of genetic structure between Sicilian and southern Italian butterfly populations, which are separated by only 3 km of ocean. They reveal how intrinsic species traits and extrinsic environmental constraints explain this pattern, demonstrating an important role for wind. Moreover, the inclusion of almost 8,000 georeferenced sequences reveals that, in spite of also being present in southern Italy, almost half of Sicilian butterfly species are more closely related to populations from other parts of Europe, Asia or North Africa. We provide further discussion on the biogeographic barrier they identify, and the potential of community‐level DNA barcoding to identify processes that structure genetic variation across communities., The authors are supported by Spanish Agencia Estatal de Investigación (CGL2017‐85718‐P), co‐financed by FEDER, with additional support to B.C.E. through the iBioGen project, funded through the European Union's Horizon 2020 research and innovation programme under grant agreement No 810729. E.J.‐G. was funded by the Ministerio de Educación y Formación Profesional through the FPU PhD fellowship (FPU18/04601) and D.S. was funded by the Ministerio de Ciencia, Innovación y Universidades through an FPI PhD fellowship (PRE2018‐083230).

Recent biological analyses suggest that reductions in dispersal ability have beenkey drivers of diversification across numerous lineages. We synthesise emergingdata to highlight similarities regarding the causes and consequences of dispersalreduction across taxa and ecosystems, as well as the diverse genomic mechanismsunderpinning these shifts. Natural selection has acted on standing genetic variationwithin taxa to drive often rapid–and in some cases parallel–losses of dispersal,and ultimately speciation. Such shifts can thus represent an important nexus be-tween adaptive and neutral diversification processes, with substantial evolutionaryconsequences. Recognition of the links between these concepts that are emergingfrom differentfields, taxa and ecosystems is transforming our understanding of thefascinating role of dispersal reduction in the formation of biodiversity., J.M.W. and G.A.M. were supported by Marsden Fund contract UOO1412 (Royal Society of New Zealand). B.C.E. and P.A.B. were supported by Ministerio de Ciencia, Innovación, y Universidades (MICIU) grant CGL2017-85718-P (Agencia Estatal de Investigación/European Regional Development Fund).

BPEC is an R package for Bayesian phylogeographic and ecological clustering which allows geographical, environmental and phenotypic measurements to be combined with deoxyribonucleic acid (DNA) sequences in order to reveal geographic structuring of DNA sequence clusters consistent with migration events. DNA sequences are modelled using a collapsed version of a simplified coalescent model projected onto haplotype trees, which subsequently give rise to constrained clusterings as migrations occur. Within each cluster, a multivariate Gaussian distribution of the covariates (geographical, environmental, phenotypic) is used. Inference follows tailored reversible jump Markov chain Monte Carlo sampling so that the number of clusters (i.e., migrations) does not need to be pre-specified. A number of output plots and visualizations are provided which reflect the posterior distribution of the parameters of interest. BPEC also includes functions that create output files which can be loaded into Google Earth. The package commands are illustrated through an example dataset of the polytypic Near Eastern brown frog Rana macrocnemis analyzed using BPEC.

Dispersal ability is known to influence geographical structuring of genetic variation within species, with a direct relationship between low vagility and population genetic structure, which can potentially give rise to allopatric speciation. However, our general understanding of the relationship between dispersal ability, population differentiation and lineage diversification is limited. To address this issue, we sampled mitochondrial DNA variation within lineages of beetles and spiders across the Canary Islands to explore the relationships between dispersal ability, differentiation within lineages and diversification. We found positive relationships between population genetic structure and diversification for both beetles and spiders. Comparisons between dispersive and non-dispersive lineages revealed significant differences for both lineage differentiation and diversification. For both taxa, non-dispersive lineages had stronger population genetic structure. Genus-level endemic species richness and proxies for diversification rate within genera were higher in non-dispersive taxa for both beetles and spiders. Comparisons of average and maximum node divergences within genera suggest that species turnover may be higher in non-dispersive genera. Our results reveal a model where dispersal limitation may shape the diversity of lineages across evolutionary timescales by positively influencing intraspecific and species diversity, moderated by higher extinction rates compared to more dispersive lineages., This work was supported by projects CGL2017-85718-P (funded by MCIN/AEI/10.13039/501100011033, Spain and EDRF, EU), and PID2020-116788GB-I00 (funded by MCIN/AEI/10.13039/501100011033, Spain) awarded to B.C.E. P.A. was funded through a Junior Leader Fellowship (LCF/BQ/PR21/11840006) by “la Caixa” Foundation (ID 100010434) and from the European Union’s Horizon 2020 research and innovation programme under the Marie Skłodowska-Curie grant agreement no. 847648. D.S. was funded by the ‘Ministerio de Ciencia e Innovación’ through a FPI PhD fellowship (PRE2018-083230) and E.J.-G. was funded by the ‘Ministerio de Educación y Formación Profesional’ through an FPU PhD fellowship (FPU18/04601). The authors wish to thank the following for assistance with fieldwork and sample sorting: Rienk Apperloo, Manuel Arechavaleta, Salvador de la Cruz, Nuria Macías, Benito Pérez, Sara Ravagni, Isa Sancibrián, Nieves Zurita, Pedro Oromí, David Hernández, Irene Santos, Heriberto López, Antonio Pérez and Carmelo Andújar. We also thank Jesús Arribas for the beetle and the non-ballooner spider images that were used to illustrate figures. We are deeply grateful to Antonia Salces-Castellano for help with the implementation of analytical scripts. We would like to thank Miquel Arnedo as well as an anonymous reviewer and Associated Editor for their useful comments on a previous version of this manuscript. Fieldwork was supported by collecting permits 246285, AFF-107/17 (sigma no. 2017-00572), A/EST-034/16, RE:2349 and REUS-11227 kindly provided by the Canary Islands Government, Cabildo of Tenerife, Cabildo of La Palma, Cabildo of El Hierro, and Cabildo de La Gomera, respectively.

Disentangling the relative role of environmental filtering and spatial processes in driving metacommunity structure across mountainous regions remains challenging, as the way we quantify spatial connectivity in topographically and environmentally heterogeneous landscapes can influence our perception of which process predominates. More empirical data sets are required to account for taxon- and context-dependency, but relevant research in understudied areas is often compromised by the taxonomic impediment. Here we used haplotype-level community DNA metabarcoding, enabled by stringent filtering of amplicon sequence variants (ASVs), to characterize metacommunity structure of soil microarthropod assemblages across a mosaic of five forest habitats on the Troodos mountain range in Cyprus. We found similar β diversity patterns at ASV and species (OTU, operational taxonomic unit) levels, which pointed to a primary role of habitat filtering resulting in the existence of largely distinct metacommunities linked to different forest types. Within-habitat turnover was correlated to topoclimatic heterogeneity, again emphasizing the role of environmental filtering. However, when integrating landscape matrix information for the highly fragmented Quercus alnifolia habitat, we also detected a major role of spatial isolation determined by patch connectivity, indicating that stochastic and niche-based processes synergistically govern community assembly. Alpha diversity patterns varied between ASV and OTU levels, with OTU richness decreasing with elevation and ASV richness following a longitudinal gradient, potentially reflecting a decline of genetic diversity eastwards due to historical pressures. Our study demonstrates the utility of haplotype-level community metabarcoding for characterizing metacommunity structure of complex assemblages and improving our understanding of biodiversity dynamics across mountainous landscapes worldwide.

AIM: Habitat diversity has been linked to the diversity and structure of island communities, however, little is known about patterns and processes within habitats. Here we aim to determine the contributions of habitat type and inferred dispersal frequency to the differences in taxonomic structure between assemblages in the same island habitat. LOCATION: The Macaronesian archipelagos (Azores, Madeira, the Canary Islands and Cabo Verde). TAXON: Spiders (Araneae). METHODS: We established forest and dry habitat sites (each with five plots) on two islands per archipelago. We collected spiders using standardised sampling protocols. We tested the differences in beta diversity separately for each habitat and for each inferred category of ballooning (an aerial dispersal strategy) frequency across geographic scales through nested non-parametric permutational multivariate analyses of variance. We then tested whether ballooning and habitat influenced heterogeneity in species composition (dispersion in beta diversity) in the two habitat types. We analysed the effects of habitat and ballooning on species abundance distribution (SAD) and rarity by fitting Gambin models and evaluating the contribution of ballooning categories to SAD. RESULTS: Communities of the same archipelago and habitat were taxonomically more similar, and beta diversity increased with geographic scale, being greater in dry habitats. There was greater species replacement among assemblages in dry habitats than in forests, with greater differences for rare ballooners. There were no differences in SAD between habitats although dry habitat sites seemed to harbour more species with low abundances (rare species) than forests. MAIN CONCLUSIONS: Habitat type does not only condition the differences between spider assemblages of the same habitat but also the scale at which they occur. These differences may be determined by the heterogeneity in the physical structure of each habitat as well as how much this structure facilitates aerial dispersal (ballooning), and should be considered in theories/hypotheses on island community assembly as well as in conservation strategies. The Government of the Canary Islands, Grant/Award Number: SE-12/02, SE-12/03 and SE-12/04; Fundação para a Ciência e a Tecnologia, Grant/Award Number: FCT-PTDC/BIABIC/0054/2014, DL57/2016/CP1375/CT0003 and UIDB/00329/2020-2024; FEDER, Grant/Award Number: 0003/2011 info:eu-repo/semantics/publishedVersion

[Aim]: Quantifying abundance distributions is critical for understanding both how communities assemble, and how community structure varies through time and space, yet estimating abundances requires considerable investment in fieldwork. Community-level population genetic data potentially offer a powerful way to indirectly infer richness, abundance and the history of accumulation of biodiversity within a community. Here we introduce a joint model linking neutral community assembly and comparative phylogeography to generate both community-level richness, abundance and genetic variation under a neutral model, capturing both equilibrium and non-equilibrium dynamics, [Location]: Global, [Methods]: Our model combines a forward-time individual-based community assembly process with a rescaled backward-time neutral coalescent model of multi-taxa population genetics. We explore general dynamics of genetic and abundance-based summary statistics and use approximate Bayesian computation (ABC) to estimate parameters underlying the model of island community assembly. Finally, we demonstrate two applications of the model using community-scale mtDNA sequence data and densely sampled abundances of an arachnid community on La Réunion. First, we use genetic data alone to estimate a summary of the abundance distribution, ground-truthing this against the observed abundances. Then, we jointly use the observed genetic data and abundances to estimate the proximity of the community to equilibrium, [Results]: Simulation experiments of our ABC procedure demonstrate that coupling abundance with genetic data leads to improved accuracy and precision of model parameter estimates compared with using abundance-only data. We further demonstrate reasonable precision and accuracy in estimating a metric underlying the shape of the abundance distribution, temporal progress towards local equilibrium and several key parameters of the community assembly process. For the insular arachnid assemblage, we find the joint distribution of genetic diversity and abundance approaches equilibrium expectations, and that the Shannon entropy of the observed abundances can be estimated using genetic data alone, [Main conclusions]: The framework that we present unifies neutral community assembly and comparative phylogeography to characterize the community-level distribution of both abundance and genetic variation through time, providing a resource that should greatly enhance understanding of both the processes structuring ecological communities and the associated aggregate demographic histories., Funding was provided by grants from FAPESP (BIOTA, 2013/50297‐0 to M.J.H. and AC Carnaval), NASA through the Dimensions of Biodiversity Program (DOB 1343578) and the National Science Foundation (DEB‐1253710 to M.J.H.). This work would not have been possible without help from the City University of New York High Performance Computing Center, with support from the National Science Foundation (CNS‐0855217 and CNS‐0958379).

Metazoan metabarcoding is emerging as an essential strategy for inventorying biodiversity,with diverse projects currently generating massive quantities of community-level data. The potential for integrating across such data sets offers new opportunities to better understand biodiversity and how it might respond to global change. However, large-scale synthesesmay be compromised if metabarcoding workflows differ from each other. There are ongoing efforts to improve standardization for the reporting of inventory data. However, harmonization at the stage of generating metabarcode data has yet to be addressed. A modular framework for harmonized data generation offers a pathway to navigate the complex structure of terrestrial metazoan biodiversity. Here, through our collective expertise as practitioners,method developers, and researchers leading metabarcoding initiatives to inventory terrestrial biodiversity, we seek to initiate a harmonized framework for metabarcode data generation, with a terrestrial arthropod module. We develop an initial set of submodules covering the 5 main steps of metabarcode data generation: (i) sample acquisition; (ii) sample processing; (iii) DNA extraction; (iv) polymerase chain reaction amplification, library preparation, and sequencing; and (v) DNA sequence and metadata deposition, providing a backbone for a terrestrial arthropod module. To achieve this, we (i) identified key points for harmonization, (ii) reviewed the current state of the art, and (iii) distilled existing knowledge within submodules, thus promoting best practice by providing guidelines and recommendations to reduce the universe of methodological options.We advocate the adoption and further development of the terrestrial arthropodmodule.We further encourage the development of modules for other biodiversity fractions as an essential step toward large-scale biodiversity synthesis through harmonization., The working group “Toward Harmonisation for the Generation of Metabarcoding Data: Soil Biodiversity and Terrestrial Arthropod modules” held in November 2020 (online) was organized by the iBioGen project, which has received funding from the European Union’s Horizon 2020 research and innovation program under grant agreement No. 810729. P.A. was funded through a Junior Leader Fellowship (LCF/BQ/PR21/11840006) by “la Caixa” Foundation (ID 100010434) and the European Union’s Horizon 2020 research and innovation program under the Marie Skłodowska-Curie grant agreement No. 847648.

Disentangling the relative role of environmental filtering and dispersal limitation in driving metacommunity structure across mountainous regions remains challenging, as the way we quantify spatial connectivity in topographically and environmentally heterogeneous landscapes can influence our perception of which process predominates. More empirical datasets are required to account for taxon- and context-dependency but relevant research is often compromised by coarse taxonomic resolution. We here employed haplotype-level community DNA metabarcoding, enabled by stringent filtering of Amplicon Sequence Variants (ASVs), to characterize metacommunity structure of soil microarthropod assemblages across a mosaic of five forest habitats on the Troodos mountain range in Cyprus. We found similar β diversity patterns at ASV and species (OTU, Operational Taxonomic Unit) levels, which pointed to a primary role of habitat filtering resulting in the existence of largely distinct metacommunities linked to different forest types. Within-habitat turnover was correlated to topoclimatic heterogeneity, again emphasizing the role of environmental filtering. However, when integrating landscape matrix information for the highly fragmented Golden Oak habitat, we also detected a major role of dispersal limitation imposed by patch connectivity, indicating that stochastic and niche-based processes synergistically govern community assembly. Alpha diversity patterns varied between ASV and OTU levels, with OTU richness decreasing with elevation and ASV richness following a longitudinal gradient, potentially reflecting a decline of genetic diversity eastwards due to historical pressures. Our study demonstrates the utility of haplotype-level community metabarcoding for characterising metacommunity structure of complex assemblages and improving our understanding of biodiversity dynamics across mountainous landscapes worldwide.

Metabarcoding of DNA extracted from community samples of whole organisms (whole organism community DNA, wocDNA) is increasingly being applied to terrestrial, marine and freshwater metazoan communities to provide rapid, accurate and high resolution data for novel molecular ecology research. The growth of this field has been accompanied by considerable development that builds on microbial metabarcoding methods to develop appropriate and efficient sampling and laboratory protocols for whole organism metazoan communities. However, considerably less attention has focused on ensuring bioinformatic methods are adapted and applied comprehensively in wocDNA metabarcoding. In this study we examined over 600 papers and identified 111 studies that performed COI metabarcoding of wocDNA. We then systematically reviewed the bioinformatic methods employed by these papers to identify the state-of-the-art. Our results show that the increasing use of wocDNA COI metabarcoding for metazoan diversity is characterised by a clear absence of bioinformatic harmonisation, and the temporal trends show little change in this situation. The reviewed literature showed (i) high heterogeneity across pipelines, tasks and tools used, (ii) limited or no adaptation of bioinformatic procedures to the nature of the COI fragment, and (iii) a worrying underreporting of tasks, software and parameters. Based upon these findings we propose a set of recommendations that we think the wocDNA metabarcoding community should consider to ensure that bioinformatic methods are appropriate, comprehensive and comparable. We believe that adhering to these recommendations will improve the long-term integrative potential of wocDNA COI metabarcoding for biodiversity science.

The genus Baezia Alonso-Zarazaga & García, 1999 is endemic to the Canary Islands, where four species were known to date. Based on morphological evidence, three new species of Baezia are described in this study: Baezia aranfaybo García & López, sp. nov. from El Hierro island, and Baezia madai García & Oromí sp. nov. and Baezia tizziri García & Andújar, sp. nov. from La Palma island. Notes on their biology, habitat, and distribution are presented. The number of taxa in this endemic Canarian genus increases to seven eyeless species. One species has been reported from the soil (endogean environment), with the other six associated with caves and the mesovoid shallow substratum (hypogean or subterranean environment). Frequent association with the presence of roots suggests that species of Baezia may inhabit the continuum represented by the endogean and hypogean environments. Identification key to the seven species are provided., This study was partly supported by the Spanish Ministry of Science (MINECO) (CGL2015-74178-JIN and CGL2017-85718-P).

Dispersal limitation has been recurrently suggested to shape both macroecological patterns and microevolutionary processes within invertebrates. However, because of potential interactions among biological, environmental, temporal, and spatial variables, causal links among flight-related traits, diversification and spatial patterns of community assembly remain elusive. Integrating genetic variation within species across whole insect assemblages, within a simplified spatial and environmental framework, can be used to reduce the impact of these potentially confounding variables. Here, we used standardized sampling and mitochondrial DNA sequencing for a whole-community characterization of the beetle fauna inhabiting a singular forested habitat (laurel forest) within an oceanic archipelago setting (Canary Islands). The spatial structure of species assemblages together with species-level genetic diversity was compared at the archipelago and island scales for 104 winged and 110 wingless beetle lineages. We found that wingless beetle lineages have: (i) smaller range sizes at the archipelago scale, (ii) lower representation in younger island communities, (iii) stronger population genetic structure, and (iv) greater spatial structuring of species assemblages between and within islands. Our results reveal that dispersal limitation is a fundamental trait driving diversity patterns at multiple hierarchical levels by promoting spatial diversification and affecting the spatial configuration of entire assemblages at both island and archipelago scales.

Biodiversity accumulates hierarchically by means of ecological and evolutionary processes and feedbacks. Within ecological communities drift, dispersal, speciation, and selection operate simultaneously to shape patterns of biodiversity. Reconciling the relative importance of these is hindered by current models and inference methods, which tend to focus on a subset of processes and their resulting predictions. Here we introduce Massive Eco-evolutionary Synthesis Simulations (MESS), a unified mechanistic model of community assembly, rooted in classic island biogeography theory, which makes temporally explicit joint predictions across three biodiversity data axes: i) species richness and abundances; ii) population genetic diversities; and iii) trait variation in a phylogenetic context. Using simulations we demonstrate that each data axis captures information at different timescales, and that integrating these axes enables discriminating among previously unidentifiable community assembly models. MESS is unique in generating predictions of community-scale genetic diversity, and in characterizing joint patterns of genetic diversity, abundance, and trait values. MESS unlocks the full potential for investigation of biodiversity processes using multi-dimensional community data including a genetic component, such as might be produced by contemporary eDNA or metabarcoding studies. We combine with supervised machine learning to fit the parameters of the model to real data and infer processes underlying how biodiversity accumulates, using communities of tropical trees, arthropods, and gastropods as case studies that span a range of data availability scenarios, and spatial and taxonomic scales., Funding was provided by grants from FAPESP (BIOTA, 2013/50297-0 to MJH and AC Carnaval), the Synthesis Centre of iDiv (DFG FZT 118), NASA through the Dimensions of Biodiversity Program (DOB 1343578) and the National Science Foundation (DEB-1253710 to MJH; DEB 1745562 to AC Carnaval; DBI 1927319 to AJR). IO was supported by the Mina Rees Dissertation Fellowship in the Sciences provided by the Graduate Center of the City University of New York. MR was supported by the Bioinformatics and Computational Biology Fellowship through the Institute for Bioinformatics and Evolutionary Studies at the University of Idaho. AJR was supported by the Santa Fe Institute Omidyar Fellowship. JR was supported by fellowships from the Natural Environment Research Council (NERC) (NE/I021179, NE/L011611/1). RSE was supported by an NWO-VICI grant. This work is a contribution to Imperial College’s Grand Challenges in Ecosystems and the Environment initiative, through JR.

Oceanic island archipelagos provide excellent models to understand evolutionary processes. Colonization events and gene flow can interact with selection to shape genetic variation at different spatial scales. Landscape-scale variation in biotic and abiotic factors may drive fine-scale selection within islands, while long-term evolutionary processes may drive divergence between distantly related populations. Here, we examine patterns of population history and selection between recently diverged populations of the Berthelot's pipit (Anthus berthelotii), a passerine endemic to three North Atlantic archipelagos. First, we use demographic trees and f3 statistics to show that genome-wide divergence across the species range is largely shaped by colonization and bottlenecks, with evidence of very weak gene flow between populations. Then, using a genome scan approach, we identify signatures of divergent selection within archipelagos at single nucleotide polymorphisms (SNPs) in genes potentially associated with craniofacial development and DNA repair. We did not detect within-archipelago selection at the same SNPs as were detected previously at broader spatial scales between archipelagos, but did identify signatures of selection at loci associated with similar biological functions. These findings suggest that similar ecological factors may repeatedly drive selection between recently separated populations, as well as at broad spatial scales across varied landscapes., This work was supported by Natural Environment Research Council (NERC) studentships to C.A.M. and C.A. (NE/L002582/1), a Norwich Research Park Science Links grant to D.S.R., M.C. and L.G.S., and a BBSRC fellowship (BB/N011759/1) and British Ecological Society Large Research Grant to L.G.S.We thank two anonymous reviewers whose insight and helpful suggestions have improved this manuscript. We thank Matthew Clark and Lawrence Percival-Alwyn for assistance generating the Berthelot's pipit genome, and Helen Hipperson, Clemens Küpper and Terry Burke (NERC Biomolecular Analysis Facility—Sheffield) for assistance with RAD sequencing. We also thank the Spanish Environment Ministry, the Canary government and the Natural Park of Madeira who gave permission for us to perform the sampling work, local governments in the Canary Islands and Madeira who provided accommodation, and the Portuguese Navy who provided transport. The research presented in this paper was carried out on the High Performance Computing Cluster supported by the Research and Specialist Computing Support service at the University of East Anglia.

Aim: To (a) evaluate if colonization of newly emerged high-elevation habitats occurred from neighbouring highlands (allopatric divergence and niche conservatism) or from within the same mountain (peripatric or parapatric divergence and niche shift), and (b) test for the effect of past landscape configurations in geographically structuring biodiversity (a West-East structure spanning the central part of the Transmexican Volcanic Belt).Location: Transmexican Volcanic Belt, Mexico. Taxa: Coleoptera and Araneae. Methods: Coleoptera and Araneae communities were collected in a comparative sampling framework. Seven sky-islands were sampled within two high-elevation habitats separated by altitude. Samples were individually sequenced for a region of the mtDNA COI gene to delimit Operational Taxonomic Units (OTUs) and haplotypes within them. Richness was estimated at the community level, as well as genetic diversity within and among mountains and habitats. The West East structure was tested with a null model and an analysis of variance with permutations. Results: Few OTUs were shared between the Abies forests and alpine grasslands. High beta diversity was found across mountains. Beta diversity between the West – East grouping is higher than expected by chance. Main conclusions: Our results indicate that colonization from other mountains followed by allopatric divergence dominates as a source of OTU formation, and that despite some probable connectivity during glacial periods existed, ancient communities and genetic structures persist., We would like to thank CONACyT for the financial support granted through a master’s scholarship to AU and for the project Ciencia Básica (178245) granted to DP. BCE is supported by Spanish Agencia Estatal de Investigación (CGL2017-85718-P), co-financed by FEDER.

Metabarcoding of Metazoa using mitochondrial genes may be confounded by both the accumulation of PCR and sequencing artefacts and the co-amplification of nuclear mitochondrial pseudogenes (NUMTs). The application of read abundance thresholds and denoising methods is efficient in reducing noise accompanying authentic mitochondrial amplicon sequence variants (ASVs). However, these procedures do not fully account for the complex nature of concomitant sequences and the highly variable DNA contribution of specimens in a metabarcoding sample. We propose, as a complement to denoising, the metabarcoding Multidimensional Abundance Threshold Evaluation (metaMATE) framework, a novel approach that allows comprehensive examination of multiple dimensions of abundance filtering and the evaluation of the prevalence of unwanted concomitant sequences in denoised metabarcoding datasets. metaMATE requires a denoised set of ASVs as input, and designates a subset of ASVs as being either authentic (mitochondrial DNA haplotypes) or nonauthentic ASVs (NUMTs and erroneous sequences) by comparison to external reference data and by analysing nucleotide substitution patterns. metaMATE (i) facilitates the application of read abundance filtering strategies, which are structured with regard to sequence library and phylogeny and applied for a range of increasing abundance threshold values, and (ii) evaluates their performance by quantifying the prevalence of nonauthentic ASVs and the collateral effects on the removal of authentic ASVs. The output from metaMATE facilitates decision-making about required filtering stringency and can be used to improve the reliability of intraspecific genetic information derived from metabarcode data. The framework is implemented in the metaMATE software (available at )., C. A. was supported by the Spanish Ministry of Economy and Competitiveness (MINECO, Spain) (CGL2015‐74178‐JIN) and Fundación CajaCanarias/Obra social “la Caixa.” B. C. E. was supported by the project CGL2017‐85718‐P (AEI, Spain/FEDER, EU). P. A., T. J. C., B. C. E. and A. P. V. were supported by the iBioGen project funded by the H2020 European Research Council, Grant/Award Number: 810729. We extend our gratitude to the regional governments of Andalucía and Canarias (Spain) for facilitating collecting of samples, to Jesús Arribas for assistance with field sampling, and Carlos Martínez for the mathematical advice.

The most remarkable feature of our planet is the diversity of its life forms, ranging from viruses and nanobacteria to blue whales and giant sequoias to satanic leaf‐tailed geckos and leafy seadragons (look them up!). Life is found in essentially all environments on earth, and the number of species living on our planet is many times greater than we could have imagined a century ago. A well‐regarded estimate pegs the number of eukaryotic species on earth at 8.7 million (±1.3 million), of which fewer than 15% are currently described (Mora et al., 2011). The diversity of prokaryotes is less clear (and highly controversial), but an analysis of 1.6 billion 16S ribosomal RNA sequences estimated that 0.8–1.6 million prokaryotic operational taxonomic units exist globally (Louca et al., 2019). While we do not know how many species are currently extant, or have existed in the past, we do know that this biodiversity is valuable, providing food, fibre and medicine, furnishing ecosystem services such as water and air purification, nutrient cycling, pollination and carbon uptake, and contributing to technological innovations ranging from biotechnology to robotics to material science. Moreover, biodiversity underlies the cultural identity of human populations and is important to human health and well‐being. Geographically, species richness increases from the Polar Regions to the tropics in terrestrial and surface marine ecosystems. Thus, some countries, especially those in tropical and subtropical regions, are endowed with much greater biodiversity than others. Unfortunately, benefits arising from the access and utilization of this biodiversity have been unequally shared, with (paradoxically) biodiversity‐poor countries often accruing the lion's share of economic gains. There can be imbalances within countries as well, wherein some segments of the population obtain greater economic benefits from biodiversity and associated traditional knowledge than indigenous peoples. The “Nagoya Protocol on Access to Genetic Resources and the Fair and Equitable Sharing of Benefits Arising from their Utilization,” which came into force in 2014, is an international agreement designed to ensure that the benefits arising from biodiversity are shared equitably (https://www.cbd.int/abs/). However, few scientific journals require compliance with the Nagoya Protocol or the reporting of benefits from biodiversity research. In this editorial, we (the editors of Molecular Ecology and Molecular Ecology Resources) express support for the Nagoya Protocol and the principle of benefit sharing. We believe that scientific journals publishing research on biodiversity can play an important role in implementing the Nagoya Protocol and in reporting on benefits generated from such research. Below, we provide background on the Nagoya Protocol, discuss the kinds of benefits that may arise from biodiversity research, describe the rationale for reporting on these benefits and introduce changes to the journals’ Data Accessibility Statements to incorporate the requirements and goals of the Nagoya Protocol.

Montane cloud forests are areas of high endemism, and are one of the more vulnerable terrestrial ecosystems to climate change. Thus, understanding how they both contribute to the generation of biodiversity, and will respond to ongoing climate change, are important and related challenges. The widely accepted model for montane cloud forest dynamics involves upslope forcing of their range limits with global climate warming. However, limited climate data provides some support for an alternative model, where range limits are forced downslope with climate warming. Testing between these two models is challenging, due to the inherent limitations of climate and pollen records. We overcome this with an alternative source of historical information, testing between competing model predictions using genomic data and demographic analyses for a species of beetle tightly associated to an oceanic island cloud forest. Results unequivocally support the alternative model: populations that were isolated at higher elevation peaks during the Last Glacial Maximum are now in contact and hybridizing at lower elevations. Our results suggest that genomic data are a rich source of information to further understand how montane cloud forest biodiversity originates, and how it is likely to be impacted by ongoing climate change., This work was financed by the Spanish Agencia Estatal de Investigación (CGL2017‐85718‐P), awarded to BCE, and co‐financed by FEDER. It was also supported by the Spanish Ministerio de Ciencia, Innovación y Universidades (EQC2018‐004418‐P), awarded to BCE. AS‐C was funded by the Spanish Ministerio de Ciencia, Innovación y Universidades through an FPU PhD fellowship (FPU014/02948). The authors thank Instituto Tecnológico y de Energías Renovables (ITER), S.A for providing access to the Teide High‐Performance Computing facility (Teide‐HPC). Fieldwork was supported by collecting permit AFF 107/17 (sigma number 2017‐00572) kindly provided by the Cabildo of Tenerife.

© 2020 by the authors., Phylogenetic relatedness is a key diversity measure for the analysis and understanding of how species and communities evolve across time and space. Understanding the nonrandom loss of species with respect to phylogeny is also essential for better-informed conservation decisions. However, several factors are known to influence phylogenetic reconstruction and, ultimately, phylogenetic diversity metrics. In this study, we empirically tested how some of these factors (topological constraint, taxon sampling, genetic markers and calibration) affect phylogenetic resolution and uncertainty. We built a densely sampled, species-level phylogenetic tree for spiders, combining Sanger sequencing of species from local communities of two biogeographical regions (Iberian Peninsula and Macaronesia) with a taxon-rich backbone matrix of Genbank sequences and a topological constraint derived from recent phylogenomic studies. The resulting tree constitutes the most complete spider phylogeny to date, both in terms of terminals and background information, and may serve as a standard reference for the analysis of phylogenetic diversity patterns at the community level. We then used this tree to investigate how partial data affect phylogenetic reconstruction, phylogenetic diversity estimates and their rankings, and, ultimately, the ecological processes inferred for each community. We found that the incorporation of a single slowly evolving marker (28S) to the DNA barcode sequences from local communities, had the highest impact on tree topology, closely followed by the use of a backbone matrix. The increase in missing data resulting from combining partial sequences from local communities only had a moderate impact on the resulting trees, similar to the difference observed when using topological constraints. Our study further revealed substantial differences in both the phylogenetic structure and diversity rankings of the analyzed communities estimated from the different phylogenetic treatments, especially when using non-ultrametric trees (phylograms) instead of time-stamped trees (chronograms). Finally, we provide some recommendations on reconstructing phylogenetic trees to infer phylogenetic diversity within ecological studies., This study was supported by the project BIODIV ISLAND-CONT (Biodiversity drivers on islands and continents–706482) funded by Marie Sklodowska-Curie Individual Fellowships (H2020-MSCA-IF-2015) to the first author N.M.-H. The research was additionally funded by four other projects that provided the material collected: 1. ERA-Net Net-Biome research framework, financed through Portuguese FCT-NETBIOME grant 0003/2011 (P.A.V.B.). 2. ERA-Net Net-Biome financed through Canary Islands Government ACIISI grants SE-12/02, SE-12/03, SE-12/04 (B.C.E.), co-financed by FEDER. 3. FCT MACDIV–FCT-PTDC/BIABIC/0054/2014 (P.A.V.B.,B.C.E., P.C., I.R.A., F.R., O.S.P., A.V.). 4. Organismo Autónomo de Parques Nacionales Spain (OAPN #485/2012) (M.A.A.). Additional support was provided by grant 2017SGR83 from the Catalan Government to M.A.A. M.D. was supported by an APIF PhD fellowship from the University of Barcelona. J.L.-F. was supported by a Juan de la Cierva Fellowship (FJCI-2015-23723). I.R.A. was funded by Portuguese funds through FCT–Fundação para a Ciência e a Tecnologia, I.P., under the Norma Transitória–DL57/2016/CP1375/CT0003. Open access funding provided by University of Helsinki.

Soil mesofauna communities are hyperdiverse and critical for ecosystem functioning. However, our knowledge on spatial structure and underlying processes of community assembly for soil arthropods is scarce, hampered by limited empirical data on species diversity and turnover. We implement a high-throughput-sequencing approach to generate comparative data for thousands of arthropods at three hierarchical levels: genetic, species and supra-specific lineages. A joint analysis of the spatial arrangement across these levels can reveal the predominant processes driving the variation in biological assemblages at the local scale. This multi-hierarchical approach was performed using haplotype-level-COI metabarcoding of entire communities of mites, springtails and beetles from three Iberian mountain regions. Tens of thousands of specimens were extracted from deep and superficial soil layers and produced comparative phylogeographic data for >1000 co-distributed species and nearly 3000 haplotypes. Local assemblages were highly distinctive between grasslands and forests, and within each of them showed strong spatial structures and high endemicity at the scale of a few kilometres or less. The local distance-decay patterns were self-similar for the haplotypes and higher hierarchical entities, and this fractal structure was very similar in all three regions, pointing to a significant role of dispersal limitation driving the local-scale community assembly. Our results from whole-community metabarcoding provide unprecedented insight into how dispersal limitations constrain mesofauna community structure within local spatial settings over evolutionary timescales. If generalized across wider areas, the high turnover and endemicity in the soil locally may indicate extremely high richness globally, challenging our current estimations of total arthropod-diversity on Earth.

Background There is an increasing demand for databases including species trait information for biodiversity and community ecology studies. The existence of trait databases is useful for comparative studies within taxa or geographical regions, but there is low availability of databases for certain organisms. Here we present an open access functional trait database for spiders from Macaronesia and the Iberian Peninsula, recording several morphological and ecological traits related to the species life histories, microhabitat and trophic preferences. New information We present a database that includes 12 biological traits for 506 spider species present in natural forests of the Iberian Peninsula (Spain) and three Macaronesian archipelagoes (Azores, Madeira and Canary Islands). The functional trait database consists of two sections: individual-level data for six morphological traits (total body size, prosoma length, prosoma width, prosoma height, tibia I length and fang length), based on direct measurements of 2844 specimens of all spider species; and species-level aggregate data for 12 traits (same 6 morphological traits as in the previous section plus dispersal ability, vertical stratification, circadian activity, foraging strategy, trophic specialization and colonization status), based on either the average of the direct measurements or bibliographic searches. This functional trait database will serve as a data standard for currently ongoing analyses that require trait and functional diversity statistics., This study was supported by the project BIODIV ISLAND-CONT (Biodiversity drivers on islands and continents – 706482) funded by Marie Sklodowska-Curie Individual Fellowships (H2020-MSCA-IF-2015) to the first author NMH. Additional support was provided by grant 2017SGR83 from the Catalan Government to MA. PB and PC are supported by AZORESBIOPORTAL–PORBIOTA (ACORES-01-0145-FEDER-000072) for the development of Functional Traits for Azorean arthropods. MD is supported by an APIF PhD fellowship from the University of Barcelona. We also thank Rutger Vos and Martin Ramirez for their valuable comments that greatly helped to improve the quality of this paper.

Geographic isolation substantially contributes to species endemism on oceanic islands when speciation involves the colonisation of a new island. However, less is understood about the drivers of speciation within islands. What is lacking is a general understanding of the geographic scale of gene flow limitation within islands, and thus the spatial scale and drivers of geographical speciation within insular contexts. Using a community of beetle species, we show that when dispersal ability and climate tolerance are restricted, microclimatic variation over distances of only a few kilometres can maintain strong geographic isolation extending back several millions of years. Further to this, we demonstrate congruent diversification with gene flow across species, mediated by Quaternary climate oscillations that have facilitated a dynamic of isolation and secondary contact. The unprecedented scale of parallel species responses to a common environmental driver for evolutionary change has profound consequences for understanding past and future species responses to climate variation., This research was supported by Spanish MINECO grants CGL2013‐42589‐P and CGL2017‐85718‐P, co‐financed by FEDER and awarded to B.C.E, and by the ERA‐Net Net‐Biome research framework, financed through Canary Island Government ACIISI grants SE‐12/04 and SE‐12/02, co‐financed by FEDER and awarded to P.O. and B.C.E respectively. A.S.‐C. was funded by the Ministerio de Educación y Formación Profesional through the FPU PhD fellowship (FPU014/02948). J.P. was funded by the MINECO through the Juan de la Cierva Program ‐ Incorporation (IJCI‐2014‐19691) and Ramón y Cajal Program (RYC‐2016‐20506), and Marie Sklodowska‐Curie COFUND, Researchers' Night and Individual Fellowships Global (MSCA grant agreement No 747238, 'UNISLAND').