BUSCADOR RECOLECTA

Predictions of plant responses to global warming frequently ignore biotic interactions and intraspecific variation across geographical ranges. Benefactor species play an important role in plant communities by protecting other taxa from harsh environments, but the combined effects of warming and beneficiary species on their performance have been largely unexamined. We analyzed the joint effects of elevated temperature and neighbor removal on the benefactor plant Silene acaulis, in factorial experiments near its low- and high-latitude range limits in Europe. We recorded growth, probability of reproduction and fruit set during 3 years. The effects of enhanced temperature were positive near the northern limit and negative in the south for some performance measures. This pattern was stronger in the presence of neighbors, possibly due to differential thermal tolerances between S. acaulis and beneficiary species in each location. Neighbors generally had a negative or null impact on S. acaulis, in agreement with previous reviews of overall effects of plant–plant interactions on benefactors. However, small S. acaulis individuals in the north showed higher growth when surrounded by neighbors. Finally, the local habitat within each location influenced some effects of experimental treatments. Overall, we show that plant responses to rising temperatures may strongly depend on their position within the geographic range, and on species interactions. Our results also highlight the need to consider features of the interacting taxa, such as whether they are benefactor species, as well as local-scale environmental variation, to predict the joint effects of global warming and biotic interactions on species and communities., This project was funded by the Swedish Research Council to W. F. Morris (Ref: 2012-42619-94710-26) and by the Spanish Ministry of Economy, Industry and Competitiveness to M. B. García (CGL2017-90040-R).

Mountains shelter high biological diversity and constitute important barriers for species distributions. They often contain sets of species whose populations occur at their range limit (peripheral species), which according to the “Centre-Periphery” hypothesis are expected to perform worse and be more vulnerable than in central positions. Our study investigates this hypothesis by examining the potential vulnerability (abundance and ecological characteristics) of the flora of the Pyrenees, a major biogeographical crossroad containing a large proportion of the total European plant diversity. We also assess the contribution of peripheral plants in lists of conservation. We compared regional area of occupancy, local abundance, elevation, habitat and soil type preferences, of more than 2,600 central, peripheral and endemic native vascular plants of the Pyrenees mountains. Their conservation status was also assessed at different spatial scales. A quarter of Pyrenean species are at their distributional limit. Like endemics, peripherals have lower continental and regional occupancy than central ones, but their local abundance does not differ significantly. Endemics and peripherals are also more likely to be soil specialists at high elevation, mainly in (sub)alpine grasslands and rocky areas. Although occurring in different ecological conditions, peripheral species at their rear-edge (mainly Boreoalpine and Eurosiberian) tend to be more widespread regionally but equally abundant locally than leading- edge species (mainly Mediterranean). Peripheral taxa constitute a large portion of Pyrenean species protected at different geographic or administrative scales (31-56%), of highest importance in the Pyrenean red list. Peripheral species show contrasting ecology at the leading- and rear-edge, contribute substantially to the high plant diversity of the Pyrenean biogeographical crossroad and to the lists of priority species for conservation. Integrative biogeographical assessments of the rarity and ecology of mountain floras provide a better overview than administrative ones to establish priorities for conservation., OPCC-POCTEFA EFA (235/1) and VULBIMON (CGL2017-90040-R)., Peer reviewed

Population sizes, threats, and abundance change through time, for monitored plants in the Aragon Region (Spain) between 2010 and 2019., Regional Government of Aragón, the European Project RESECOM (LIFE+12 NAT/ES/000180), the OAPN (DYNBIO, grant 1656/2015), the Research Spanish Agency (VULBIMON, grant CGL2017-90040-R), and the Diputación Provincial de Huesca (2019), Peer reviewed

[Description of methods used for collection/generation of data] We built a genus-level phylogeny using the workflow proposed by Roquet et al. (2013). We downloaded from Genbank three conserved chloroplastic regions (rbcL, matK and ndhF) plus the ITS region for a subset of families, which we aligned separately by taxonomic clustering.

[Methods for processing the data] We aligned all coding sequence clusters with MACSE (Ranwez et al. 2011) and non-coding ones with MAFFT (Katoh and Standlye 2013), and trimmed all alignments with TrimAl (Capella-Gutiérrez et al. 2009). We concatenated all alignments to obtain a supermatrix. We then conducted maximum-likelihood (ML) phylogenetic inference analyses with RAxML (Stamatakis 2014), applying the most appropriate partitioning scheme and substitution model obtained with PartitionFinder (Lanfear et al. 2012) and a supertree constraint at the family-level obtained with the online software Phylomatic v.3 (tree R20120829). Specifically, we performed 100 independent tree searches and selected the best ML tree (the one with the highest probability).

[Instrument- or software-specific information needed to interpret/reproduce the data] Any software for manipulation or analysis of phylogenies such as R packages ape or picante.

[Standards and calibration information, if appropriate] The best ML tree was dated applying the penalized likelihood method in treePL (Smith and O'Meara,2012) and the following node calibrations: we fixed the node corresponding to the ancestor of eudicots at 125 Ma based on the earliest eudicot fossil (Hughes and McDougall 1990), and applied minimum age constraints to 15 nodes based on fossil information extracted from Smith and Beaulieu (2010) and Bell et al. (2010).

5. Environmental/experimental conditions:, Agencia Estatal de Investigación, Project VULVIMON (Reference: CGL2017-90040-R)., Peer reviewed

Biodiversity loss is one of the most relevant consequences of climate change. Therefore, identifying areas and environmental features that allow certain organisms to be less exposed to the effects of the current global warming is priority for biodiversity conservation. In this study, we describe a novel approach for the identification of microclimatic refugia in rugged mountain areas, specifically for the detection of most thermally stable areas, using an unmanned aerial vehicle (UAV) capable of recording in the visible and thermal infrared spectral bands. We estimated land surface temperatures (LST) at very-high spatial resolution in six topographically complex sectors of the Pyrenees (NE Spain), across seasons with vegetative activity (summer 2020, autumn 2020, spring 2021, and summer 2021), and at two thermally contrasted times of the day (early in the morning: LSTmin, and in the afternoon: LSTmax). LST were validated with a network of miniaturized temperature sensors in the field. LSTmin and LSTmax allowed us to calculate the daily thermal range of each sector across the seasons, and thus the most thermally stable areas over the year. To reveal the importance of different variables on low and narrow thermal ranges we applied Gradient Boosted Models to seven terrain variables derived from ALS-LiDAR (slope, northness, eastness, heat load, wind exposure index, SAGA's topographic wetness index, and vector ruggedness measure) and a proxy of forest density through the three-dimensional point clouds of the UAV data. The northness was the variable that most promoted thermal stability, followed by the slope and forest density, so that microclimatic refugia resulted to be located in northern slopes, small sites under rocky cliffs, and forested areas. Our results demonstrate that thermal UAVs can become promising tools for the identification of microclimatic refugia in topographically complex areas, providing information at unprecedented spatial resolution, and thus of high interest for biodiversity conservation., This work was supported by the Spanish Association of Terrestrial Ecology (AEET) through the program “Grants for research projects led by young researchers 2019”; the Spanish Ministry of Science, Innovation, and Universities through an FPU predoctoral contract granted to R.H. (FPU18/05027); and the Spanish Research Agency through the VULBIMON (CGL2017-90040-R) and REFUGIA (PID2021-129056OB-I00) projects to M.B.G., Peer reviewed

© The Author(s) 2022. Open Access This article is licensed under a Creative Commons Attribution 4.0 International License, which permits use, sharing, adaptation, distribution and reproduction in any medium or format, as long as you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons licence, and indicate if changes were made. The images or other third party material in this article are included in the article's Creative Commons licence, unless indicated otherwise in a credit line to the material. If material is not included in the article's Creative Commons licence and your intended use is not permitted by statutory regulation or exceeds the permitted use, you will need to obtain permission directly from the copyright holder. To view a copy of this licence, visit http://creativecommons.org/licenses/by/4.0/., Climatic refugia have often been associated with hotspots of richness and endemism, and identified on the basis of molecular or paleobotanical information. Here, we apply a phylogenetic analysis to 18,000 plant communities distributed across the Pyrenees, a south European mountain range, to identify climatic refugia from imprints of relictuality inferred from species’ evolutionary distinctiveness (ED). We produced a genus-level phylogenetic tree to calculate the standardized mean ED value of plant communities (cED). Then, we explored which habitats concentrate the plant communities with the highest cED and the interrelated effect of past (long-term climatic stability) and present (topographic and spatial position) factors. Results show strong differences of cED among habitats: forests ranked first, followed by some open habitats like high altitude wetlands. Climate stability and roughness positively influenced cED. A weak negative association resulted between the two diversity measurements (richness and endemism rate) and also with cED. We propose that forests acted as “mobile refugia” during the glacial-interglacial periods, supported by paleoenvironmental reconstructions revealing continuous presence at regional scale of key broadleaved trees at that time. Azonal habitats like the endemic-poor humid communities at high elevation would have also played an important role as more permanent microrefugia. Our approach identifies a variety of habitats and plant assemblages that have successfully withstood past climate change in different ways, and therefore would hold an important evolutionary potential to cope with current climate change. Given their potential role in preserving biodiversity, they should be integrated in future conservation agendas., Open Access funding provided thanks to the CRUE-CSIC agreement with Springer Nature. This study is a product of the VULBIMON project (CGL2017-90040-R), and had the support of the PYCACHU project (PID2019-106050RB-I00). HM received a Grant from the Spanish Ministry of Science (PRE2018-083868)., Peer reviewed

© 2022 The Authors. Diversity and Distributions published by John Wiley & Sons Ltd. This is an open access article under the terms of the Creative Commons Attribution License, which permits use, distribution and reproduction in any medium, provided the original work is properly cited., [Aim] Mountains shelter high biological diversity and constitute both important barriers and confluence areas for species. They often contain species whose populations occur at their range limit (peripheral species), which according to the “Centre-Periphery” hypothesis (CPH) are expected to occur in marginal environments, exhibit low abundance and consequently high vulnerability. Our study investigates this hypothesis for the flora of the Pyrenees, a biogeographical crossroads containing a large proportion of the total European plant diversity., [Location] Pyrenees., [Methods] We determined whether more than 2600 native plant species were endemic to the Pyrenees or found at the centre or periphery of their whole distribution range within the mountain chain. We then compared the ecological preferences, local and regional abundance, and conservation status among central, peripheral and endemic species., [Results] A quarter of the flora was found at its geographic range limit within the Pyrenees. Endemic and peripheral species were more likely to be soil specialists in alpine grasslands and rocks, and exhibited smaller regional ranges than central species, but their local abundance did not tend to differ. Peripheral species at their southern range edge were more widespread regionally than at their northern range edge. Peripheral taxa were more prevalent in the Pyrenean red list of threatened species (55%) compared to national and regional protection lists (40% and 31%, respectively)., [Main conclusions] Peripheral species contribute substantially to the diversity of the Pyrenean flora. They follow the predictions of the CPH given their occurrence in scarce habitats, their low regional abundance and their high vulnerability according to the Pyrenean red list, although they tend to show similar local abundances as other species and are infrequent in protection lists. Integrative and cross-border assessments of the ecology and rarity of mountain floras provide better estimates of their vulnerability and ensure more efficient prioritization of their conservation., Species records were compiled through the European projects FLORAPYR and FLORAPYR AVANCE, funded by the Pyrenean Work Community, a cross-border cooperation consortium. The authors thank all regional project leaders and multiple Pyrenean collaborators involved in the project, for gathering and putting together much information used here (http://www.atlasflorapyrenaea.eu/src/home/index.php?idma=0), and the OPCC-POCTEFA (EFA322/19 EFA 235/11). SP and MBG received funding by the Spanish Ministry of Science and Technology through a doctoral grant (FPI, BES-2011-045169), the project CAMBIO (CGL2010-21642) and VULBIMON (CGL2017-90040-R)., Peer reviewed

Little is known about how much continuous landscape transformation might affect the most vulnerable elements of biodiversity. In this study, we quantified changes in the normalized difference vegetation index (NDVI) over the past 35 years across locations with threatened plants and in Natura 2000 (N2000) protected areas, in an environmentally heterogeneous region of Southern Europe. First, we estimated the intensity and duration of NDVI gains and losses based on Landsat time series using the LandTrendr algorithm in Google Earth Engine. Then, we tested: 1) whether populations of threatened plants were located in more stable sites than non-threatened plants (i.e., lower NDVI changes); 2) whether NDVI changes around populations of threatened plants differed across habitats and inside/outside N2000 areas, and 3) whether lower NDVI changes occurred in N2000 areas than unprotected areas, thereby indicating their effectiveness at preserving biodiversity. Threatened plants tended to be concentrated in sites with less change irrespective of the habitat where they occurred and their location within protected areas. Occurrence in stable sites also reduced the risk associated with small-sized populations. N2000 areas were in line with the overall greening trend but they experienced less loss events than the unprotected areas, thereby supporting their role in slowing down human-induced land cover changes. Our approach demonstrates how long-term remote sensing monitoring can help to assess the effects of both slow processes and drastic landscape transformation events on priority plants in a comprehensive and rapid manner. This method can identify hidden patterns in extensive regions and guide effective conservation management., This study was funded by the grant JAEINT18_EX_0379, VULBIMON project (ref. CGL2017-90040-R), and eLTER PLUS project (ID: 871128)., Peer reviewed

© 2022 The Authors. Oikos published by John Wiley & Sons Ltd on behalf of Nordic Society Oikos. This is an open access article under the terms of the Creative Commons Attribution License, which permits use, distribution and reproduction in any medium, provided the original work is properly cited., Species rarity has puzzled ecologists for decades owing to its theoretical fundamentals and practical implications. The main concern that rarity raises from a conservation standpoint is that rare species tend to have higher extinction risk. Understanding the underlying phylogenetic patterns of rarity can shed light on the evolutionary processes yielding rarity, contribute to forecasting potential vulnerable species or lineages, and assess the consequences of the loss of rare species. Here, we apply Rabinowitz's rarity classification scheme and explore phylogenetic patterns in the flora of the Pyrenees, a temperate mountain range. We first categorized species according to their geographic range (endemicity and regional geographic range), habitat specialization and local abundance. Then we analyzed the phylogenetic signal of each rarity type, along with the phylogenetic association between species according to their rarity, and decomposed the variation of rarity among taxonomic levels to better understand its diversification through time. Finally, we estimated the expected loss of phylogenetic diversity (PD) in the regional flora if rare species go extinct. All rarity types showed a certain degree of phylogenetic signal, with endemics, species of limited regional geographic range and habitat specialists forming groups of phylogenetically close species. The decomposition of variation supported these results, as the rarity types with the strongest signal had more variation explained by higher taxonomic levels. The loss of these groups of species had different consequences based on the type of rarity, with the disappearance of habitat specialists having the strongest, negative effect on PD. Our results show a general pattern of phylogenetic association in rarity among the flora of the Pyrenees, which leads to increased vulnerability along certain branches of the Tree of Life. Phylogenetic patterns should be taken into account in conservation planning to effectively protect all facets of biodiversity., This study is a product of the VULBIMON project (CGL2017-90040-R). HMC received a grant from the Spanish Ministry of Science (grant no. PRE2018-083868)., Peer reviewed

In the present scenario of climatic change, climatic refugia will be of paramount importance for species per- sistence. Topography can generate a considerable climatic heterogeneity over short distances, which is often disregarded in macroclimatic predictive models. Here we investigate the role of rocky habitats as microclimatic refugia by combining two different analyses: exploring a thermal mechanism whereby rocky habitats might serve as refugia, and examining if the biogeographic pattern shows a high abundance of relict, endemic and peripheral species.
The thermal profile of two populations of relict and endemic plant species occurring in Pyrenean cliffs was investigated by infrared images and in situ temperature data-loggers. Despite occurring in crevices of a south oriented slope, Androsace cylindrica showed a narrower daily range of temperature than the surrounding matrix, thereby avoiding extreme high temperatures. Borderea chouardii, of tropical ancestors, also occurred in patches where temperatures were buffered during the growth season, experiencing lower mean temperatures than the surrounding matrix and nearby areas during the warmer part of the day, and similar temperatures during the colder. The rocky habitats of both species, therefore, reduced temperature ranges and exposition to extreme climatic events. Compared to other habitats, the rocky ones also harboured a very high fraction of both endemics and peripheral plant populations according to the largest vegetation dataset available in the Pyrenees (18,800 plant inventories and 400,000 records). Our results suggest an association between the habitats of relicts, en- demics and species at their distribution limit, driven by a stabilizing effect of rocky habitats on extreme tem- peratures. Given the important role of rocky habitats as hotspots of singular and unique plants, their char- acterization seems a sensible first step to identify potential refugia in the context of climate change.

Biodiversity loss is one of the most relevant consequences of climate change. Therefore, identifying areas and environmental features that allow certain organisms to be less exposed to the effects of the current global warming is priority for biodiversity conservation. In this study, we describe a novel approach for the identification of microclimatic refugia in rugged mountain areas, specifically for the detection of most thermally stable areas, using an unmanned aerial vehicle (UAV) capable of recording in the visible and thermal infrared spectral bands. We estimated land surface temperatures (LST) at very-high spatial resolution in six topographically complex sectors of the Pyrenees (NE Spain), across seasons with vegetative activity (summer 2020, autumn 2020, spring 2021, and summer 2021), and at two thermally contrasted times of the day (early in the morning: LSTmin, and in the afternoon: LSTmax). LST were validated with a network of miniaturized temperature sensors in the field. LSTmin and LSTmax allowed us to calculate the daily thermal range of each sector across the seasons, and thus the most thermally stable areas over the year. To reveal the importance of different variables on low and narrow thermal ranges we applied Gradient Boosted Models to seven terrain variables derived from ALS-LiDAR (slope, northness, eastness, heat load, wind exposure index, SAGA's topographic wetness index, and vector ruggedness measure) and a proxy of forest density through the three-dimensional point clouds of the UAV data. The northness was the variable that most promoted thermal stability, followed by the slope and forest density, so that microclimatic refugia resulted to be located in northern slopes, small sites under rocky cliffs, and forested areas. Our results demonstrate that thermal UAVs can become promising tools for the identification of microclimatic refugia in topographically complex areas, providing information at unprecedented spatial resolution, and thus of high interest for biodiversity conservation.

Research in global change ecology relies heavily on global climatic grids derived from estimates of air temperature in open areas at around 2 m above the ground. These climatic grids do not reflect conditions below vegetation canopies and near the ground surface, where critical ecosystem functions occur and most terrestrial species reside. Here, we provide global maps of soil temperature and bioclimatic variables at a 1-km resolution for 0-5 and 5-15 cm soil depth. These maps were created by calculating the difference (i.e. offset) between in situ soil temperature measurements, based on time series from over 1200 1-km pixels (summarized from 8519 unique temperature sensors) across all the world's major terrestrial biomes, and coarse-grained air temperature estimates from ERA5-Land (an atmospheric reanalysis by the European Centre for Medium-Range Weather Forecasts). We show that mean annual soil temperature differs markedly from the corresponding gridded air temperature, by up to 10°C (mean = 3.0 ± 2.1°C), with substantial variation across biomes and seasons. Over the year, soils in cold and/or dry biomes are substantially warmer (+3.6 ± 2.3°C) than gridded air temperature, whereas soils in warm and humid environments are on average slightly cooler (−0.7 ± 2.3°C). The observed substantial and biome-specific offsets emphasize that the projected impacts of climate and climate change on near-surface biodiversity and ecosystem functioning are inaccurately assessed when air rather than soil temperature is used, especially in cold environments. The global soil-related bioclimatic variables provided here are an important step forward for any application in ecology and related disciplines. Nevertheless, we highlight the need to fill remaining geographic gaps by collecting more in situ measurements of microclimate conditions to further enhance the spatiotemporal resolution of global soil temperature products for ecological applications.