Volcanic tephra deposition dataset based on interpolated field measurements following the 2021 Tajogaite Eruption on La Palma, Canary Islands, Spain.

In 2021, the Tajogaite Volcano erupted along the western slope of the Cumbre Vieja on the island of La Palma, Canary Islands, Spain. Volcanic tephra blanketed a substantial proportion of the island. By our estimations, approximately 23,000,000 m3 of pyroclastic ashes and more coarse-grained particles were deposited unto La Palma's land surface in addition to the lava flow. Five months following the initial eruption, we measured the depth of the new ash layer across the island. We combined this data with drone-based observations to compile a dataset comprising the point distribution of ash depth. A spatial interpolation was then performed using Inverse Distance Weighting (IDW) to estimate the ash depth across the island at a 2 m spatial resolution. The interpolation performed well, yielding a root mean squared error (RMSE) value of 0.34 and thus, the dataset offers immense reuse potential for spatial inquiries related to evolutionary traits, vegetation patterns, and vegetation response to disturbance on oceanic islands. In addition, the data can be used to test different spatial interpolation techniques in an effort to improve the accuracy achieved using IDW.

Diurnal temperature range as a key predictor of plants' elevation ranges globally.

A prominent hypothesis in ecology is that larger species ranges are found in more variable climates because species develop broader environmental tolerances, predicting a positive range size-temperature variability relationship. However, this overlooks the extreme temperatures that variable climates impose on species, with upper or lower thermal limits more likely to be exceeded. Accordingly, we propose the 'temperature range squeeze' hypothesis, predicting a negative range size-temperature variability relationship. We test these contrasting predictions by relating 88,000 elevation range sizes of vascular plants in 44 mountains to short- and long-term temperature variation. Consistent with our hypothesis, we find that species' range size is negatively correlated with diurnal temperature range. Accurate predictions of short-term temperature variation will become increasingly important for extinction risk assessment in the future.

A multitaxonomic assessment of Natura 2000 effectiveness across European biogeographic regions.

The Natura 2000 (N2K) protected area (PA) network is a crucial tool to limit biodiversity loss in Europe. Despite covering 18% of the European Union's (EU) land area, its effectiveness at conserving biodiversity across taxa and biogeographic regions remains uncertain. Testing this effectiveness is, however, difficult because it requires considering the nonrandom location of PAs, and many possible confounding factors. We used propensity score matching and accounted for the confounding effects of biogeographic regions, terrain ruggedness, and land cover to assess the effectiveness of N2K PAs on the distribution of 1769 species of conservation priority in the EU's Birds and Habitats Directives, including mammals, birds, amphibians, reptiles, arthropods, fishes, mollusks, and vascular and nonvascular plants. We compared alpha, beta, and gamma diversity between matched selections of protected and unprotected areas across EU's biogeographic regions with generalized linear models, generalized mixed models, and nonparametric tests for paired samples, respectively, for each taxonomic group and for the entire set of species. PAs in N2K hosted significantly more priority species than unprotected land, but this difference was not consistent across biogeographic regions or taxa. Total alpha diversity and alpha diversity of amphibians, arthropods, birds, mammals, and vascular plants were significantly higher inside PAs than outside, except in the Boreal biogeographical region. Beta diversity was in general significantly higher inside N2K PAs than outside. Similarly, gamma diversity had the highest values inside PAs, with some exceptions in Boreal and Atlantic regions. The planned expansion of the N2K network, as dictated by the European Biodiversity Strategy for 2030, should therefore target areas in the southern part of the Boreal region where species diversity of amphibians, arthropods, birds, mammals, and vascular plants is high and species are currently underrepresented in N2K.

Análisis multitaxonómico de la efectividad de Natura 2000 en las regiones biogeográficas de Europa Resumen La red de áreas protegidas (AP) de Natura 2000 (N2K) es una herramienta importante para reducir la pérdida de biodiversidad en Europa. A pesar de que cubre el 18% del área terrestre de la UE, todavía es incierta la efectividad que tiene para conservar la biodiversidad en los taxones y las regiones biogeográficas. Sin embargo, es complicado analizar esta efectividad porque requiere considerar la ubicación no azarosa de las AP y la posibilidad de muchos factores confusos. Usamos el pareamiento por puntaje de propensión y consideramos los efectos confusos de las regiones biogeográficas, lo accidentado del terreno y la cobertura del suelo para analizar la efectividad de las AP de N2K en la distribución de 1,769 especies (mamíferos, aves, anfibios, reptiles, artrópodos, peces, moluscos y plantas vasculares y no vasculares) con prioridad de conservación en las Directivas de Aves y Hábitats de la UE. Comparamos la diversidad alfa, beta y gamma entre las selecciones pareadas de las áreas protegidas y no protegidas en las regiones biogeográficas de la UE con los modelos generalizados lineales, mixtos y pruebas no paramétricas de las muestras pareadas, respectivamente, para cada grupo taxonómico y para el conjunto completo de especies. Las áreas protegidas en N2K tuvieron una mayoría significativa de especies prioritarias en comparación con el suelo no protegido, pero esta diferencia no fue coherente entre los taxones y las regiones biogeográficas. La diversidad alfa total y la diversidad alfa de anfibios, artrópodos, aves, mamíferos y plantas vasculares fue significativamente mayor dentro de las AP que fuera de ellas, excepto en la región biogeográfica boreal. La diversidad beta fue significativamente más alta dentro de las AP de N2K que fuera de ellas. De forma similar, la diversidad gamma tuvo los valores más altos dentro de las AP, salvo algunas excepciones en las regiones boreal y atlántica. Por lo tanto, la expansión planeada de la red N2K, como dicta la Estrategia de la UE sobre Biodiversidad para 2030, debería enfocarse en las áreas del sur de la región boreal, donde es alta la diversidad de especies de anfibios, artrópodos, aves, mamíferos y plantas vasculares y cuyas especies están poco representadas dentro de N2K.

Long-term evidence of differential resistance and resilience of grassland ecosystems to extreme climate events.

Grassland ecosystems are affected by the increasing frequency and intensity of extreme climate events (e.g., droughts). Understanding how grassland ecosystems maintain their functioning, resistance, and resilience under climatic perturbations is a topic of current concern. Resistance is the capacity of an ecosystem to withstand change against extreme climate, while resilience is the ability of an ecosystem to return to its original state after a perturbation. Using the growing season Normalized Difference Vegetation Index (NDVIgs, an index of vegetation growth) and the Standardized Precipitation Evapotranspiration Index (a drought index), we evaluated the response, resistance, and resilience of vegetation to climatic conditions for alpine grassland, grass-dominated steppe, hay meadow, arid steppe, and semi-arid steppe in northern China for the period 1982-2012. The results show that NDVIgs varied significantly across these grasslands, with the highest (lowest) NDVIgs values in alpine grassland (semi-arid steppe). We found increasing trends of greenness in alpine grassland, grass-dominated steppe, and hay meadow, while there were no detectable changes of NDVIgs in arid and semi-arid steppes. NDVIgs decreased with increasing dryness from extreme wet to extreme dry. Alpine and steppe grasslands exhibited higher resistance to and lower resilience after extreme wet, while lower resistance to and higher resilience after extreme dry conditions. No significant differences in resistance and resilience of hay meadow under climatic conditions suggest the stability of this grassland under climatic perturbations. This study concludes that highly resistant grasslands under conditions of water surplus are low resilient, but low resistant ecosystems under conditions of water shortage are highly resilient.

Long-term trajectories of non-native vegetation on islands globally.

Human-mediated changes in island vegetation are, among others, largely caused by the introduction and establishment of non-native species. However, data on past changes in non-native plant species abundance that predate historical documentation and censuses are scarce. Islands are among the few places where we can track human arrival in natural systems allowing us to reveal changes in vegetation dynamics with the arrival of non-native species. We matched fossil pollen data with botanical status information (native, non-native), and quantified the timing, trajectories and magnitude of non-native plant vegetational change on 29 islands over the past 5000 years. We recorded a proportional increase in pollen of non-native plant taxa within the last 1000 years. Individual island trajectories are context-dependent and linked to island settlement histories. Our data show that non-native plant introductions have a longer and more dynamic history than is generally recognized, with critical implications for biodiversity baselines and invasion biology.

Links to rare climates do not translate into distinct traits for island endemics.

Current models of island biogeography treat endemic and non-endemic species as if they were functionally equivalent, focussing primarily on species richness. Thus, the functional composition of island biotas in relation to island biogeographical variables remains largely unknown. Using plant trait data (plant height, leaf area and flower length) for 895 native species in the Canary Islands, we related functional trait distinctiveness and climate rarity for endemic and non-endemic species and island ages. Endemics showed a link to climatically rare conditions that is consistent with island geological change through time. However, functional trait distinctiveness did not differ between endemics and non-endemics and remained constant with island age. Thus, there is no obvious link between trait distinctiveness and occupancy of rare climates, at least for the traits measured here, suggesting that treating endemic and non-endemic species as functionally equivalent in island biogeography is not fundamentally wrong.

Challenges and solutions to biodiversity conservation in arid lands.

The strategic goals of the United Nations and the Aichi Targets for biodiversity conservation have not been met. Instead, biodiversity has continued to rapidly decrease, especially in developing countries. Setting a new global biodiversity framework requires clarifying future priorities and strategies to bridge challenges and provide representative solutions. Hyper-arid, arid, and semi-arid lands (herein, arid lands) form about one third of the Earth's terrestrial surface. Arid lands contain unique biological and cultural diversity, and biodiversity loss in arid lands can have a disproportionate impact on these ecosystems due to low redundancy and a high risk of trophic cascades. They contain unique biological and cultural diversity and host many endemic species, including wild relatives of key crop plants. Yet extensive agriculture, unsustainable use, and global climate change are causing an irrecoverable damage to arid lands, with far-reaching consequences to the species, ground-water resources, ecosystem productivity, and ultimately the communities' dependant on these systems. However, adequate research and effective policies to protect arid land biodiversity and sustainability are lacking because a large proportion of arid areas are in developing countries, and the unique diversity in these systems is frequently overlooked. Developing new priorities for global arid lands and mechanisms to prevent unsustainable development must become part of public discourse and form the basis for conservation efforts. The current situation demands the combined efforts of researchers, practitioners, policymakers, and local communities to adopt a socio-ecological approach for achieving sustainable development (SDGs) in arid lands. Applying these initiatives globally is imperative to conserve arid lands biodiversity and the critical ecological services they provide for future generations. This perspective provides a framework for conserving biodiversity in arid lands for all stakeholders that will have a tangible impact on sustainable development, nature, and human well-being.

The fate of terrestrial biodiversity during an oceanic island volcanic eruption.

Volcanic activity provides a unique opportunity to study the ecological responses of organisms to catastrophic environmental destruction as an essential driver of biodiversity change on islands. However, despite this great scientific interest, no study of the biodiversity at an erupting volcano has yet been undertaken. On La Palma (Canary archipelago), we quantified the main species affected and their fate during the 85-day eruption (September-December 2021). Our main objective consisted of monitoring the biodiversity subjected to critical stress during this volcanic eruption. We found that all biodiversity within a 2.5 km radius was severely affected after the first two weeks. It is challenging to assess whether volcanism can drive evolutionary traits of insular organisms. Examples are the adaptation of an endemic conifer to high temperatures, selection of functional plant types-secondary woodiness-, effects of the disappearance of invertebrates and their influence in trophic nets and vertebrate trophic plasticity. However, our data suggest that such previous evolutionary changes might continue to favour their resilience during this eruption. Lastly, it is a very good opportunity to assess the extent to which these periodic volcanic catastrophes may constitute temporary windows of repeated opportunities for the evolution and speciation of oceanic island biota.

The Spectral Species Concept in Living Color.

Biodiversity monitoring is an almost inconceivable challenge at the scale of the entire Earth. The current (and soon to be flown) generation of spaceborne and airborne optical sensors (i.e., imaging spectrometers) can collect detailed information at unprecedented spatial, temporal, and spectral resolutions. These new data streams are preceded by a revolution in modeling and analytics that can utilize the richness of these datasets to measure a wide range of plant traits, community composition, and ecosystem functions. At the heart of this framework for monitoring plant biodiversity is the idea of remotely identifying species by making use of the 'spectral species' concept. In theory, the spectral species concept can be defined as a species characterized by a unique spectral signature and thus remotely detectable within pixel units of a spectral image. In reality, depending on spatial resolution, pixels may contain several species which renders species-specific assignment of spectral information more challenging. The aim of this paper is to review the spectral species concept and relate it to underlying ecological principles, while also discussing the complexities, challenges and opportunities to apply this concept given current and future scientific advances in remote sensing.

Biodiversity showed positive effects on resistance but mixed effects on resilience to climatic extremes in a long-term grassland experiment.

Understanding the role of biodiversity in maintaining ecosystem functioning and stability under increasing frequency and magnitude of climatic extremes has fascinated ecologists for decades. Although growing evidence suggests that biodiversity affects ecosystem productivity and buffers ecosystem against climatic extremes, it remains unclear whether the stability of an ecosystem is caused by its resistance against disturbances or resilience towards perturbations or both. In attempting to explore how species richness affects resistance and resilience of above-ground net primary productivity (ANPP) against climatic extremes, we analyzed the grassland ANPP of the long-running (1997-2020) Bayreuth Biodiversity experiment in Germany. We used the Standardized Precipitation Evapotranspiration Index to identify climatic conditions based on 5- and 7-class classifications of climatic conditions. Mixed-effects models and post-hoc test show that ANPP varied significantly among different intensities (e.g. moderate or extreme) and directions (e.g. dry or wet) of climatic conditions, with the highest ANPP in extreme wet and the lowest in extreme dry conditions. Resistance and resilience of ANPP to climatic extremes in different intensities were examined by linear-mixed effects models and we found that species richness increased ecosystem resistance against all dry and wet climatic extremes, but decreased ecosystem resilience towards all dry climatic extremes. Species richness had no effects on ecosystem resilience towards wet climatic extremes. When the five level of species richness treatment (i.e., 1, 2, 4, 8, and 16 species) were considered, the relationships between species richness and resistance and resilience of ANPP under extreme wet and dry conditions remained similar. Our study emphasizes that plant communities with greater species richness need to be maintained to stabilize ecosystem productivity and increase resistance against different climatic extremes.

Ericaceous vegetation of the Bale Mountains of Ethiopia will prevail in the face of climate change.

Climate change impacts the structure, functioning, and distribution of species and ecosystems. It will shift ecosystem boundaries, potentially affecting vulnerable ecosystems, such as tropical Africa's high mountain ecosystems, i.e., afroalpine ecosystems, and their highly susceptible uniquely adapted species. However, ecosystems along these mountains are not expected to respond similarly to the change. The ericaceous woody vegetation, located between the low-elevation broadleaf forests and high-elevation afroalpine vegetation, are anticipated to be affected differently. We hypothesize that projected climate change will result in an upward expansion and increasing dominance of ericaceous vegetation, which will negatively impact the endemic rich afroalpine ecosystems of the extensive Sanetti plateau. Hence, we modeled the impact of future climate change on the distribution of ericaceous vegetation and discussed its effect on bordering ecosystems in the Bale Mountains. We applied four familiar correlative modeling approaches: bioclim, domain, generalized linear methods, and support vector machines. We used WorldClim's bioclimatic variables as environmental predictors and two representative concentration pathways (RCPs) of the IPCC Fifth Assessment Report climate change scenarios, namely RCP4.5 and RCP8.5 for future climate projection. The results indicate increased ericaceous vegetation cover on the midaltitude of northwestern and northern parts of the massif, and the Sanetti plateau. We observed upward range expansion and increase of close ericaceous vegetation in midaltitudes, while receding from the lower range across the massif. Moreover, the current ericaceous vegetation range correlates to the temperature and precipitation trends, reaffirming the critical role of temperature and precipitation in determining species distributions along elevational gradients. The results indicate the high likelihood of considerable changes in this biodiversity hotspot in Eastern Africa.

Landscape fragmentation of the Natura 2000 network and its surrounding areas.

Habitat loss from anthropogenic development has led to an unprecedented decline in global biodiversity. Protected areas (PAs) exist to counteract this degradation of ecosystems. In the European Union, the Natura 2000 (N2k) network is the basis for continent-wide conservation efforts. N2k is the world's largest coordinated network of protected areas. However, threats to ecosystems do not stop at the borders of PAs. As measured by a landscape fragmentation metric, anthropogenic development can affect the interiors of PAs. To ensure the long-term viability of the N2k network of PAs, this paper attempts to quantify the degree to which N2k sites are insulated from development pressures. We use a comprehensive dataset of effective mesh density (seff) to measure aggregate fragmentation inside and within a 5 km buffer surrounding N2k sites. Our results show a strong correlation (R² = 0.78) between fragmentation (seff) within and around N2k sites. This result applies to all biogeographical regions in Europe. Only a narrow majority (58.5%) of N2k sites are less fragmented than their surroundings. Remote and mountainous regions in northern Europe, the Alps, parts of Spain, and parts of eastern Europe show the lowest levels of fragmentation. These regions tend to hold the largest N2k sites as measured by area. In contrast, central and western Europe show the highest fragmentation levels within and around N2k sites. 24.5% of all N2k sites are classified as highly to very-highly fragmented. N2k PA age since initial protection does not correlate with the difference in exterior and interior fragmentation of N2k PAs. These results indicate that PAs in Europe are not sheltered from anthropogenic pressures leading to fragmentation. Hence, we argue that there is a high potential for improving PA efficacy by taking pre-emptive action against encroaching anthropogenic fragmentation and by targeting scarce financial resources where fragmentation pressures can be mitigated through enforced construction bans inside PAs.

Vegetation plot and trait data from phonolitic and basaltic rocks on La Palma (Canary Islands, Spain).

Geodiversity promotes biodiversity by increasing habitat heterogeneity. In times of a global biodiversity decline, data about diversity on such geological elements gains importance, also regarding conservation and restoration. In the Canary Islands, phonolitic rocks are geological elements of volcanic origin that represent additional habitat for species. In the southern part of the island La Palma, phonolite rocks can be encountered in matrices of young lava. We provide biotic and abiotic records from 60 plots of 2 m × 2 m, sampled on phonolitic and neighbouring basaltic outcrops at four different sites that cover a wide environmental gradient. The recorded parameters were species abundances (percentage cover), plant individuals' frequency (number of plant individuals per plot and species), plant growth height (in cm), and canopy diameter (in cm). Additionally, lichen cover (percentage cover) was estimated. To describe site specific characteristics, we recorded plot surface dynamics (3D rock surface), aspect, and inclination. This data set covers vegetation and trait data comparing phonolites and basalts on La Palma. It can be used for island biogeography, vegetation ecology and conservation sciences to help support fundamental research related to insular biodiversity and endemism, and to identify priority areas for protection and preservation in the Canary Islands.

Chikungunya Beyond the Tropics: Where and When Do We Expect Disease Transmission in Europe?

Chikungunya virus disease (chikungunya) is a mosquito-borne infectious disease reported in at least 50 countries, mostly in the tropics. It has spread around the globe within the last two decades, with local outbreaks in Europe. The vector mosquito Aedes albopictus (Diptera, Culicidae) has already widely established itself in southern Europe and is spreading towards central parts of the continent. Public health authorities and policymakers need to be informed about where and when a chikungunya transmission is likely to take place. Here, we adapted a previously published global ecological niche model (ENM) by including only non-tropical chikungunya occurrence records and selecting bioclimatic variables that can reflect the temperate and sub-tropical conditions in Europe with greater accuracy. Additionally, we applied an epidemiological model to capture the temporal outbreak risk of chikungunya in six selected European cities. Overall, the non-tropical ENM captures all the previous outbreaks in Europe, whereas the global ENM had underestimated the risk. Highly suitable areas are more widespread than previously assumed. They are found in coastal areas of the Mediterranean Sea, in the western part of the Iberian Peninsula, and in Atlantic coastal areas of France. Under a worst-case scenario, even large areas of western Germany and the Benelux states are considered potential areas of transmission. For the six selected European cities, June-September (the 22th-38th week) is the most vulnerable time period, with the maximum continuous duration of a possible transmission period lasting up to 93 days (Ravenna, Italy).

Biotic homogenization destabilizes ecosystem functioning by decreasing spatial asynchrony.

Our planet is facing significant changes of biodiversity across spatial scales. Although the negative effects of local biodiversity (α diversity) loss on ecosystem stability are well documented, the consequences of biodiversity changes at larger spatial scales, in particular biotic homogenization, that is, reduced species turnover across space (ß diversity), remain poorly known. Using data from 39 grassland biodiversity experiments, we examine the effects of ß diversity on the stability of simulated landscapes while controlling for potentially confounding biotic and abiotic factors. Our results show that higher ß diversity generates more asynchronous dynamics among local communities and thereby contributes to the stability of ecosystem productivity at larger spatial scales. We further quantify the relative contributions of α and ß diversity to ecosystem stability and find a relatively stronger effect of α diversity, possibly due to the limited spatial scale of our experiments. The stabilizing effects of both α and ß diversity lead to a positive diversity-stability relationship at the landscape scale. Our findings demonstrate the destabilizing effect of biotic homogenization and suggest that biodiversity should be conserved at multiple spatial scales to maintain the stability of ecosystem functions and services.

Diversity of European habitat types is correlated with geography more than climate and human pressure.

Habitat richness, that is, the diversity of ecosystem types, is a complex, spatially explicit aspect of biodiversity, which is affected by bioclimatic, geographic, and anthropogenic variables. The distribution of habitat types is a key component for understanding broad-scale biodiversity and for developing conservation strategies. We used data on the distribution of European Union (EU) habitats to answer the following questions: (i) how do bioclimatic, geographic, and anthropogenic variables affect habitat richness? (ii) Which of those factors is the most important? (iii) How do interactions among these variables influence habitat richness and which combinations produce the strongest interactions? The distribution maps of 222 terrestrial habitat types as defined by the Natura 2000 network were used to calculate habitat richness for the 10 km × 10 km EU grid map. We then investigated how environmental variables affect habitat richness, using generalized linear models, generalized additive models, and boosted regression trees. The main factors associated with habitat richness were geographic variables, with negative relationships observed for both latitude and longitude, and a positive relationship for terrain ruggedness. Bioclimatic variables played a secondary role, with habitat richness increasing slightly with annual mean temperature and overall annual precipitation. We also found an interaction between anthropogenic variables, with the combination of increased landscape fragmentation and increased population density strongly decreasing habitat richness. This is the first attempt to disentangle spatial patterns of habitat richness at the continental scale, as a key tool for protecting biodiversity. The number of European habitats is related to geography more than climate and human pressure, reflecting a major component of biogeographical patterns similar to the drivers observed at the species level. The interaction between anthropogenic variables highlights the need for coordinated, continental-scale management plans for biodiversity conservation.

Deriving risk maps from epidemiological models of vector borne diseases: State-of-the-art and suggestions for best practice.

Epidemiological models (EMs) are widely used to predict the temporal outbreak risk of vector-borne diseases (VBDs). EMs typically use the basic reproduction number (R0), a threshold quantity, to indicate risk. To provide an overall view of the risk, these model outputs can be transformed into spatial risk maps, using various aggregation methods (e.g. average R0 over time, cumulative number of days with R0 > 1). However, there is no standardized methodology available for this. Depending on the specific aggregation methods used, the yielded spatial risk maps may have considerably different interpretations. Additionally, the method used to visualize the aggregated data also affects the perceived spatial patterns. In this review, we compare commonly used aggregation and visualization methods and discuss the respective interpretation of risk maps. Research publications using epidemiological modelling methods were drawn from Web of Science. Only publications containing maps of R0 transformed from EMs were considered for the analysis. An example EM was applied to illustrate how aggregation and visualization methods affect the final presentations of risk maps. Risk maps can be generated to show duration, intensity and spatio-temporal dynamics of potential outbreak risk of VBDs. We show that 1) different temporal aggregation methods lead to different interpretations; 2) similar spatial patterns do not necessarily bear the same meaning; 3) visualization methods considerably affect how results are perceived, and thus should be applied with caution. We recommend mapping both intensity and duration of the VBD outbreak risk, using small time-steps to show spatio-temporal dynamics when possible.

A grid-based map for the Biogeographical Regions of Europe.

BACKGROUND: Biogeographical units are widely adopted in ecological research and nature conservation management, even though biogeographical regionalisation is still under scientific debate. The European Environment Agency provided an official map of the European Biogeographical Regions (EBRs), which contains the official boundaries used in the Habitats and Birds Directives. However, these boundaries bisect cells in the official EU 10 km × 10 km grid used for many purposes, including reporting species and habitat data, meaning that 6881 cells overlap two or more regions. Therefore, superimposing the EBRs vector map over the grid creates ambiguities in associating some cells with European Biogeographical Regions. NEW INFORMATION: To provide an operational tool to unambiguously define the boundaries of the eleven European Biogeographical Regions, we provide a specifically developed raster map of Grid-Based European Biogeographical Regions (GB-EBRs). In this new map, the borders of the EBRs are reshaped to coherently match the standard European 10 km × 10 km grid imposed for reporting tasks by Article 17 of the Habitats Directive and used for many other datasets. We assign each cell to the EBR with the largest area within the cell.

Diversification in evolutionary arenas-Assessment and synthesis.

Understanding how and why rates of evolutionary diversification vary is a key issue in evolutionary biology, ecology, and biogeography. Evolutionary rates are the net result of interacting processes summarized under concepts such as adaptive radiation and evolutionary stasis. Here, we review the central concepts in the evolutionary diversification literature and synthesize these into a simple, general framework for studying rates of diversification and quantifying their underlying dynamics, which can be applied across clades and regions, and across spatial and temporal scales. Our framework describes the diversification rate (d) as a function of the abiotic environment (a), the biotic environment (b), and clade-specific phenotypes or traits (c); thus, d ~ a,b,c. We refer to the four components (a-d) and their interactions collectively as the "Evolutionary Arena." We outline analytical approaches to this framework and present a case study on conifers, for which we parameterize the general model. We also discuss three conceptual examples: the Lupinus radiation in the Andes in the context of emerging ecological opportunity and fluctuating connectivity due to climatic oscillations; oceanic island radiations in the context of island formation and erosion; and biotically driven radiations of the Mediterranean orchid genus Ophrys. The results of the conifer case study are consistent with the long-standing scenario that low competition and high rates of niche evolution promote diversification. The conceptual examples illustrate how using the synthetic Evolutionary Arena framework helps to identify and structure future directions for research on evolutionary radiations. In this way, the Evolutionary Arena framework promotes a more general understanding of variation in evolutionary rates by making quantitative results comparable between case studies, thereby allowing new syntheses of evolutionary and ecological processes to emerge.