eBryoSoil: a citizen science application to monitor changes in soil ecosystems.

Biological soil covers (BSCs) play a pivotal role in ecosystem functioning by enhancing soil stability, mediating nutrient cycling, and influencing soil hydrology. Recognized as ecosystem engineers, they can physically modify, maintain, or create habitats, facilitating plant community development. Through these intricate interactions, BSCs contribute significantly to ecological processes, highlighting their importance in the overall health and functionality of the ecosystems of the Iberian Peninsula. Here we present the results obtained from the contributions of the citizen scientists uploaded from November 2019 to January 2021 with eBryoSoil, an app that allows citizens to participate in mapping the BSC communities across the Iberian Peninsula. Here, we emphasize the importance of habitats and consequently, their interaction with climatic variables for the persistence of BSCs (lichens and bryophytes). Conservation efforts targeted at preserving diverse habitats are essential to ensure the continued presence of lichen and bryophyte communities. Despite challenges posed by the SARS-CoV-2 outbreak, this citizen science project demonstrated success in utilizing a specifically tailored app to gather valuable information on BSC communities, providing insights into their vulnerability to climate change. This program serves as an illustrative example of how citizen science can effectively identify and study vulnerable habitats, offering a blueprint for future studies focused on understudied organisms.

The interplay between climatic niche evolution, polyploidy and reproductive traits explains plant speciation in the Mediterranean Basin: a case study in Centaurium (Gentianaceae).

Speciation and diversification patterns in angiosperms are frequently shaped by niche evolution. Centaurium Hill is a Mediterranean genus with ca. 25 species, of which 60% are polyploids (tetra- and hexaploids), distributed mainly in the Mediterranean Basin and in areas with temperate and arid climates of Asia, Europe, North-Central Africa and North America. The evolutionary history of this genus has been studied using morphological, biogeographical and molecular approaches, but its climatic niche characterization and its relation with genome evolution (chromosome number and ploidy level) has not been addressed yet. Thus, this study aims to identify the role of the evolution of climatic niche, ploidy level, life cycle and floral traits in the diversification of Centaurium. Climatic niche characterization involved estimating present climate preferences using quantitative data and reconstructing ancestral niches to evaluate climatic niche shifts. The evolution of climatic niche towards selective optima determined by ploidy level (three ploidy levels) and different binary traits (polyploidy, floral size, floral display, herkogamy and life cycle) was addressed under the Ornstein-Uhlenbeck model. Chromosome number evolution was inferred using the ChromoSSE model, testing if changes are clado- or anagenetic. Chromosome number evolution and its link with cladogenesis, life cycle and floral traits was modeled on the phylogeny. The reconstruction of the ancestral niches shows that Centaurium originated in a mild climate and diversified to both humid and cold as well as to dry and warmer climates. Niche conservatism was estimated in the climatic niche of the ancestors, while the climatic niche of the current taxa experienced transitions from their ancestors' niche. Besides, the evolution of climatic niche towards multiple selective optima determined by the studied traits was supported, life cycle optima receiving the highest support. The reconstruction of chromosome number transitions shows that the rate of speciation process resulting from chromosomal changes (chromosomal cladogenesis) is similar to that of non-chromosomal cladogenesis. Additionally, dependent evolution of floral size, floral display and herkogamy with chromosome number variation was supported. In conclusion, polyploidization is a crucial process in the Mediterranean region that assisted speciation and diversification into new areas with different climates, entailing niche shifts and evolution of reproductive strategies.

Unfolding the dynamics of ecosystems undergoing alternating wet-dry transitional states.

A significant fraction of Earth's ecosystems undergoes periodic wet-dry alternating transitional states. These globally distributed water-driven transitional ecosystems, such as intermittent rivers and coastal shorelines, have traditionally been studied as two distinct entities, whereas they constitute a single, interconnected meta-ecosystem. This has resulted in a poor conceptual and empirical understanding of water-driven transitional ecosystems. Here, we develop a conceptual framework that places the temporal availability of water as the core driver of biodiversity and functional patterns of transitional ecosystems at the global scale. Biological covers (e.g., aquatic biofilms and biocrusts) serve as an excellent model system thriving in both aquatic and terrestrial states, where their succession underscores the intricate interplay between these two states. The duration, frequency, and rate of change of wet-dry cycles impose distinct plausible scenarios where different types of biological covers can occur depending on their desiccation/hydration resistance traits. This implies that the distinct eco-evolutionary potential of biological covers, represented by their trait profiles, would support different functions while maintaining similar multifunctionality levels. By embracing multiple alternating transitional states as interconnected entities, our approach can help to better understand and manage global change impacts on biodiversity and multifunctionality in water-driven transitional ecosystems, while providing new avenues for interdisciplinary studies.

Exploring the impact of data curation criteria on the observed geographical distribution of mosses.

Biodiversity data records contain inaccuracies and biases. To overcome this limitation and establish robust geographic patterns, ecologists often curate records keeping those that are most suitable for their analyses. Yet, this choice is not straightforward and the outcome of the analysis may vary due to a trade-off between data quality and volume. This problem is particularly recurrent for less-studied groups with patchy sampling effort. The latitudinal pattern of mosses richness remains inconsistent across studies and these may emerge purely from sampling artefacts. Our main objective here is to assess the effect of different curation criteria on this spatial pattern in the Temperate Northern Hemisphere (above 20° latitude). We contrasted the geographical distribution of moss species records and the latitude-species richness relation obtained under different data curation scenarios. These scenarios comprehend five sources of taxonomical standardisations and eight data cleaning filters. The analyses are based on the selection of well-surveyed cells at 100 km cell resolution. The application of some 'data curation scenarios' severely affects the number of records selected for analysis and substantially changes the proportion of richness per cell. The sensitivity to data curation becomes detectable at regional and at the cell scales showing a large shift in the latitudinal richness peak in Europe, from 60° N to 45° N latitude, when only preserved specimens are selected and duplicates based on date of collection and coordinates are excluded. Our results stress the importance of justifying the criteria used for filtering biodiversity data retrieved from biodiversity databases to avoid detecting misleading patterns. Curating records under particular criteria compromises the information in some areas displaying different spatial information of mosses. This problem can be ameliorated if data filtering is combined with identifying well-surveyed cells, render relatively constant results under different combinations of filtering even for less well-known groups such as mosses.

A Protocol to Retrieve and Curate Spatial and Climatic Data from Online Biodiversity Databases Using R.

Ecological and evolutionary studies often require high quality biodiversity data. This information is readily available through the many online databases that have compiled biodiversity data from herbaria, museums, and human observations. However, the process of preparing this information for analysis is complex and time consuming. In this study, we have developed a protocol in R language to process spatial data (download, merge, clean, and correct) and extract climatic data, using some genera of the ginseng family (Araliaceae) as an example. The protocol provides an automated way to process spatial and climatic data for numerous taxa independently and from multiple online databases. The script uses GBIF, BIEN, and WorldClim as the online data sources, but can be easily adapted to include other online databases. The script also uses genera as the sampling unit but provides a way to use species as the target. The cleaning process includes a filter to remove occurrences outside the natural range of the taxa, gardens, and other human environments, as well as erroneous locations and a spatial correction for misplaced occurrences (i.e., occurrences within a distance buffer from the coastal boundary). Additionally, each step of the protocol can be run independently. Thus, the protocol can begin with data cleaning, if the database has already been compiled, or with climatic data extraction, if the database has already been parsed. Each line of the R script is commented so that it can also be run by users with little knowledge of R.

Moss establishment success is determined by the interaction between propagule size and species identity.

Colonization of new habitat patches is a key aspect of metacommunity dynamics, particularly for sessile organisms. Mosses can establish in new patches through fragmentation, with different vegetative structures acting as propagules. Despite the importance of these propagules for successful colonization the specific aspects that favour moss colonization by vegetative propagules remain poorly understood, including the effect of propagule size. We examine the intra- and interspecific variation of establishment and colonization success in culture of propagules of different sizes in six widespread soil moss species of contrasting growth form (Dicranum scoparium, Homalothecium aureum, Hypnum cupressiforme, Ptychostomum capillare, Syntrichia ruralis and Tortella squarrosa). We obtained three different size classes of propagules from artificially fragmented vegetative material, and assessed their establishment under controlled light and temperature conditions. We characterize the size, shape, apparent viability, morphological type and size changes due to hydration states of the propagules, all of them traits with potentially significant influence in their dispersal pattern and establishment. Then we assess the effect of these traits on moss establishment, using indicators of surface establishment (number of established shoots and colonized surface) and biomass production (viable biomass) as proxies of colonization success. The establishment indicators related to colonization surface and biomass production differ among species and propagule sizes. The magnitude of the interspecific differences of all indicators of establishment success was larger at the smaller propagule size class. T. squarrosa was the most successful species, and D. scoparium showed the lowest performance. We also found interspecific differences in the hydration dynamics of the propagules. The process of establishment by vegetative fragments operates differently among moss species. Besides, differences between hydration states in propagules of some species could be part of syndromes for both dispersal and establishment. This study unveils several functional traits relevant for moss colonization, such as wet versus dry area and length of fragments, which may improve our understanding of their spatial dynamics.

Climatic niche pre-adaptation facilitated island colonization followed by budding speciation in the Madeiran ivy (Hedera maderensis, Araliaceae).

The path followed by species in the colonization of remote oceanic islands ultimately depends on their phylogenetic constraints and ecological responses. In this study, we aim to evaluate the relative role of geographical and ecological forces in the origin and evolution of the Madeiran ivy (Hedera maderensis), a single-species endemic belonging to the western polyploid clade of Hedera. To determine the phylogenetic placement of H. maderensis within the western polyploid clade, we analyzed 40 populations (92 individuals) using genotyping-by-sequencing and including Hedera helix as outgroup. Climatic niche differences among the study species were evaluated using a database with 867 records representing the entire species ranges. To test species responses to climate, 13 vegetative and reproductive functional traits were examined for 70 populations (335 individuals). Phylogenomic results revealed a nested pattern with H. maderensis embedded within the south-western Iberian H. iberica. Gradual niche differentiation from the coldest and most continental populations of H. iberica to the warm and stable coastal population sister to H. maderensis parallels the geographical pattern observed in the phylogeny. Similarity in functional traits is observed for H. maderensis and H. iberica. The two species show leaves with higher specific leaf area (SLA), lower leaf dry matter content (LDMC) and thickness and fruits with lower pulp fraction than the other western polyploid species H. hibernica. Acquisition of a Macaronesian climatic niche and the associated functional syndrome in mainland European ivies (leaves with high SLA, and low LDMC and thickness, and fruits with less pulp content) was a key step in the colonization of Madeira by the H. iberica/H. maderensis lineage, which points to climatic pre-adaptation as key in the success of island colonization (dispersal and establishment). Once in Madeira, budding speciation was driven by geographical isolation, while ecological processes are regarded as secondary forces with a putative impact in the lack of further in situ diversification.

Evaluation of tropical-temperate transitions: An example of climatic characterization in the Asian Palmate group of Araliaceae.

PREMISE: There has been a great increase in using climatic data in phylogenetic studies over the past decades. However, compiling the high-quality spatial data needed to perform accurate climatic reconstructions is time-consuming and can result in poor geographical coverage. Therefore, researchers often resort to qualitative approximations. Our aim was to evaluate the climatic characterization of the genera of the Asian Palmate Group (AsPG) of Araliaceae as an exemplar lineage of plants showing tropical-temperate transitions. METHODS: We compiled a curated worldwide spatial database of the AsPG genera and created five raster layers representing bioclimatic regionalizations of the world. Then, we crossed the database with the layers to climatically characterize the AsPG genera. RESULTS: We found large disagreement in the climatic characterization of genera among regionalizations and little support for the climatic nature of the tropical-temperate distribution of the AsPG. Both results are attributed to the complexity of delimiting tropical, subtropical, and temperate climates in the world and to the distribution of the study group in regions with transitional climatic conditions. CONCLUSIONS: The complexity in the climatic classification of this example of the tropical-temperate transitions calls for a general climatic revision of other tropical-temperate lineages. In fact, we argue that, to properly evaluate tropical-temperate transitions across the tree of life, we cannot ignore the complexity of distribution ranges.

Competition-induced transgenerational plasticity influences competitive interactions and leaf decomposition of offspring.

Phenotypic plasticity, within and across generations (transgenerational plasticity), allows organisms and their progeny to adapt to the environment without modification of the underlying DNA. Recent findings suggest that epigenetic modifications are important mediators of such plasticity. However, empirical studies have, so far, mainly focused on plasticity in response to abiotic factors, overlooking the response to competition. We tested for within-generation and transgenerational phenotypic plasticity triggered by plant-plant competition intensity, and we tested whether it was mediated via DNA methylation, using the perennial, apomictic herb Taraxacum brevicorniculatum in four coordinated experiments. We then tested the consequences of transgenerational plasticity affecting competitive interactions of the offspring and ecosystem processes, such as decomposition. We found that, by promoting differences in DNA methylation, offspring of plants under stronger competition developed faster and presented more resource-conservative phenotypes. Further, these adjustments associated with less degradable leaves, which have the potential to reduce nutrient turnover and might, in turn, favour plants with more conservative traits. Greater parental competition enhanced competitive abilities of the offspring, by triggering adaptive phenotypic plasticity, and decreased offspring leaf decomposability. Our results suggest that competition-induced transgenerational effects could promote rapid adaptations and species coexistence and feed back on biodiversity assembly and nutrient cycling.

Positive associations among rare species and their persistence in ecological assemblages.

According to the competitive exclusion principle, species with low competitive abilities should be excluded by more efficient competitors; yet, they generally remain as rare species. Here, we describe the positive and negative spatial association networks of 326 disparate assemblages, showing a general organization pattern that simultaneously supports the primacy of competition and the persistence of rare species. Abundant species monopolize negative associations in about 90% of the assemblages. On the other hand, rare species are mostly involved in positive associations, forming small network modules. Simulations suggest that positive interactions among rare species and microhabitat preferences are the most probable mechanisms underpinning this pattern and rare species persistence. The consistent results across taxa and geography suggest a general explanation for the maintenance of biodiversity in competitive environments.

BtM, a Low-cost Open-source Datalogger to Estimate the Water Content of Nonvascular Cryptogams.

Communities of nonvascular cryptogams, such as mosses or lichens, are an important part of the Earth's biodiversity, contributing to the regulation of the carbon and nitrogen cycles in many ecosystems. Being poikilohydric organisms, they do not actively control their internal water content and need a humid environment to activate their metabolism. Therefore, studying water relationships of nonvascular cryptogams is crucial to understand both their diversity patterns and their functions in the ecosystems. We present the BtM datalogger, a low-cost open-source platform for the study of the water content of nonvascular cryptogams. The datalogger is designed to measure ambient temperature, humidity, and conductance from up to eight samples simultaneously. We provide a design for a printed circuit board (PCB), a detailed protocol to assemble the components, and the required source code. All this makes the assembly of the BtM datalogger accessible to any research group, even to those without previous specialized knowledge. Therefore, the design presented here has the potential to help popularize the use of this type of device among ecologists and field biologists.

Crown damage and the mortality of tropical trees.

What causes individual tree death in tropical forests remains a major gap in our understanding of the biology of tropical trees and leads to significant uncertainty in predicting global carbon cycle dynamics. We measured individual characteristics (diameter at breast height, wood density, growth rate, crown illumination and crown form) and environmental conditions (soil fertility and habitat suitability) for 26 425 trees ≥ 10 cm diameter at breast height belonging to 416 species in a 52-ha plot in Lambir Hills National Park, Malaysia. We used structural equation models to investigate the relationships among the different factors and tree mortality. Crown form (a proxy for mechanical damage and other stresses) and prior growth were the two most important factors related to mortality. The effect of all variables on mortality (except habitat suitability) was substantially greater than expected by chance. Tree death is the result of interactions between factors, including direct and indirect effects. Crown form/damage and prior growth mediated most of the effect of tree size, wood density, fertility and habitat suitability on mortality. Large-scale assessment of crown form or status may result in improved prediction of individual tree death at the landscape scale.

Spores potentially dispersed to longer distances are more tolerant to ultraviolet radiation: A case study in the moss genus Orthotrichum.

PREMISE OF THE STUDY: Ultraviolet (UV) radiation influences the viability of algal spores and seed-plant pollen depending on the species, the dose, and the wavelength. In bryophytes, one of the dominant groups of plants in many habitats, UV radiation could determine their spore dispersal strategy, and such data are critical for reconstructing the ancestral state in plants and for determining the distribution range and persistence of bryophyte species. METHODS: Spores of four bryophyte species of the moss genus Orthotrichum that were either hygrochastic or xerochastic (spores dispersed under wet or dry conditions, respectively) were exposed to realistic doses of UV radiation under laboratory conditions. Spore viability was evaluated through germination experiments and, for the first time in bryophytes, ultrastructural observations. Given that the UV-B doses used were relatively higher than the UV-A doses, the UV effect was probably due more to UV-B than UV-A wavelengths. KEY RESULTS: All four species reduced their spore germination capacity in a UV dose-dependent manner, concomitantly increasing spore ultrastructural damage (cytoplasmic and plastid alterations). Most spores eventually died when exposed to the highest UV dose. Interestingly, spores of hygrochastic species were much more UV-sensitive than those of xerochastic species. CONCLUSIONS: UV tolerance determines moss spore viability, as indicated by germination capacity and ultrastructural damage, and differs between spores of species with different dispersal strategies. Specifically, the higher UV tolerance of xerochastic spores may enable them to be dispersed to longer distances than hygrochastic spores, thus extending more efficiently the distribution range of the corresponding species.

Shifts in the importance of the species pool and environmental controls of epiphytic bryophyte richness across multiple scales.

Species richness is influenced by a nested set of environmental factors, but how do these factors interact across several scales? Our main aim is to disentangle the relative importance of environmental filters and the species pool on the richness of epiphytic bryophytes across spatial scales. To do so, we sampled epiphytic bryophytes in 43 oak forests across the northwest of the Iberian Peninsula. As predictors we used climate, descriptors of forest structure and micro-environment. We applied structural equation modeling to relate these variables with richness and cover at three scales: locality (forest), stand (three stands per forest), and sample (a quadrate in a tree). We assumed top-down relationships, so that large-scale variables influenced lower scale variables, and in which cover directly influenced richness. Richness at the next larger scale (locality to stand and stand to sample) is considered a surrogate of the species pool and included as a predictor of richness at the next smaller scale. Environmental variables explain locality richness, but as we decrease the spatial scale, its importance decreases and the dependence on species pool increases. In addition, we found unexpected bottom-up relationships (between micro-scale environment to locality richness). Our results point to the scale dependence of niche vs. neutral processes: niche processes are important at the locality (forest) scale, while neutral processes are significant at the small (sample) scale. We propose a modified conceptualization of the factors influencing biodiversity at different spatial scales by adding links across different scales (between micro-environment and locality-scale richness in our study).

The effects of experimentally supplied lead nitrate on three common Mediterranean moss species.

We assess here, through an experimental simulation using lead nitrate, the response to lead deposition of three common Mediterranean bryophyte species in the family Pottiaceae. Five concentrations of lead nitrate (from 0 to 10-3 M) were sprayed for 4 months on plants belonging to Tortula muralis (reported as toxitolerant), Syntrichia ruralis (medium-tolerant), and Tortula subulata (less tolerant). The three species showed a remarkably high tolerance to lead nitrate, with a low incidence of damage even at concentrations as high as 10-4 M. The maximum concentration (10-3 M), although resulting eventually in serious damages in the gametophyte of the three species (high mortality rates in S. ruralis and T. subulata, or a significant percentage of damaged tissue in T. muralis), did not prevent the production of sporophytes in the two species with fertile samples (T. muralis and T. subulata). Growth parameters show limited value as bioindicators of lead deposition, as they only show clear effects at very high concentrations. Besides, we identified the existence of a lead exclusion strategy mediated by mucilage using histochemical analyses and scanning electron microscopy combined with energy-dispersive X-ray spectroscopy. This mechanism can hamper the usefulness of these mosses in quantitative estimation of lead deposition.

Does spore ultrastructure mirror different dispersal strategies in mosses? A study of seven iberian orthotrichum species.

Most mosses have xerochastic dispersal (i.e., they open their capsules when conditions are dry), which is thought to favor long-distance dispersal. However, there are several species that use a hygrochastic strategy: spores are dispersed when conditions are wet. The significance of this strategy in the Mediterranean region is unknown. In this study, we explored whether ultrastructural features related to differences in spore resistance may explain these different strategies of spore dispersal. To this end, we examined the ultrastructural features of the spores of seven closely related species in the moss genus Orthotrichum. These species all grow as epiphytes in sub-Mediterranean forests, and the group includes both xerochastic and hygrochastic members. First, we found that the spore wall layers exhibit several features previously undescribed in mosses. Second, we discovered that there are only subtle differences in spore ultrastructure with regards to spore wall thickness, the degree of plastid development, or the storage substances used. We suggest that the hygrochastic dispersal in mosses from Mediterranean environments might be related to a safe-site strategy, rather than to drought avoidance, and we underscore the necessity of conducting spore ultrastructural studies on a greater number of bryophyte species.