Microrefugia and microclimate: Unraveling decoupling potential and resistance to heatwaves.

Microrefugia, defined as small areas maintaining populations of species outside their range margins during environmental extremes, are increasingly recognized for their role in conserving species in the face of climate change. Understanding their microclimatic dynamics becomes crucial with global warming leading to severe temperature and precipitation changes. This study investigates the phenomenon of short-term climatic decoupling within microrefugia and its implications for plant persistence in the Mediterranean region of southeastern France. We focus on microrefugia's ability to climatically disconnect from macroclimatic trends, examining temperature and Vapor Pressure Deficit (VPD) dynamics in microrefugia, adjacent control plots, and weather stations. Our study encompasses both "normal" conditions and heatwave episodes to explore the role of microrefugia as thermal and moisture insulators during extreme events. Landscape attributes such as relative elevation, solar radiation, distance to streams, and vegetation height are investigated for their contribution to short-term decoupling. Our results demonstrate that microrefugia exhibit notable decoupling from macroclimatic trends. This effect is maintained during heatwaves, underscoring microrefugia's vital role in responding to climatic extremes. Importantly, microrefugia maintain lower VPD levels than their surroundings outside and during heatwaves, potentially mitigating water stress for plants. This study advances our understanding of microclimate dynamics within microrefugia and underscores their ecological importance for plant persistence in a changing climate. As heatwaves become more frequent and severe, our findings provide insights into the role of microrefugia in buffering but also decoupling against extreme climatic events and, more generally, against climate warming. This knowledge emphasizes the need to detect and protect existing microrefugia, as they can be integrated into conservation strategies and climate change adaptation plans.

Global seed dormancy patterns are driven by macroclimate but not fire regime.

Seed dormancy maximizes plant recruitment in habitats with variation in environmental suitability for seedling establishment. Yet, we still lack a comprehensive synthesis of the macroecological drivers of nondormancy and the different classes of seed dormancy: physiological dormancy, morphophysiological dormancy and physical dormancy. We examined current geographic patterns and environmental correlates of global seed dormancy variation. Combining the most updated data set on seed dormancy classes for > 10 000 species with > 4 million georeferenced species occurrences covering all of the world's biomes, we test how this distribution is driven by climate and fire regime. Seed dormancy is prevalent in seasonally cold and dry climates. Physiological dormancy occurs in relatively dry climates with high temperature seasonality (e.g. temperate grasslands). Morphophysiological dormancy is more common in forest-dominated, cold biomes with comparatively high and evenly distributed precipitation. Physical dormancy is associated with dry climates with strong seasonal temperature and precipitation fluctuations (e.g. deserts and savannas). Nondormancy is associated with stable, warm and wetter climates (e.g. tropical rain forest). Pyroclimate had no significant effect on the distribution of seed dormancy. The environmental drivers considered in this study had a comparatively low predictive power, suggesting that macroclimate is just one of several global drivers of seed dormancy.

Yellow-legged gull populations (Larus michahellis) link the history of landfills to soil eutrophication and time-related vegetation changes on small Mediterranean islands.

Seabird colonies have a strong influence on both the physical and chemical soil parameters and plant communities of the islands where they settle to nest. Scientists have studied the effects of the demographic explosion of seabird populations, but few have explored the long-term effects when the colonies were in decline. The aim of this study was to investigate diachronic changes over a 24 year period of soil parameters, floristic composition and plant functionnal types (Raunkiaer growth forms and Grime life strategies) up to the decrease of the number of nesting yellow-legged gulls (Larus michahellis Naumann, 1840) on Mediterranean islands. We used 78 permanent plots to survey the vegetation and the soil parameters on 9 islands and one mainland area within the Calanques National Park (south east of France), for three periods (i.e., 1997, 2008, 2021). Since 1997, the increase of nesting gulls has caused a nitrogen and pH increase and organic carbon and C/N ratio decrease, although the values were still higher than mainland plots without nesting gulls. This has led to changes in plant species composition e.g., higher values of N favouring the development of ruderal plant species, still present in high frequency in 2021. Furthermore, plant species highly tolerant to disturbances (i.e., R Grime strategy) in harsh environments were still favoured even after the decline of gull abundance. However, both the frequency of the chamaephytes and the vegetation cover has increased with the decline of gull colony. In 2021, measures of trace elements' concentrations and calculation of pollution load index (Cu, Pb and Zn) reveals relatively low multi-contamination levels on the mainland and the archipelagos. On naturally oligotrophic and semi-arid Mediterranean islands, gull colonies induce a persistent alteration in soil characteristics that still influences plant communities (composition and functional types), 11 years after the decline in bird abundance.

Bridging the gap between microclimate and microrefugia: A bottom-up approach reveals strong climatic and biological offsets.

In the context of global warming, a clear understanding of microrefugia-microsites enabling the survival of species populations outside their main range limits-is crucial. Several studies have identified forcing factors that are thought to favor the existence of microrefugia. However, there is a lack of evidence to conclude whether, and to what extent, the climate encountered within existing microrefugia differs from the surrounding climate. To investigate this, we adopt a "bottom-up" approach, linking marginal disconnected populations to microclimate. We used the southernmost disconnected and abyssal populations of the circumboreal herbaceous plant Oxalis acetosella in Southern France to study whether populations in sites matching the definition of "microrefugia" occur in particularly favorable climatic conditions compared to neighboring control plots located at distances of between 50 to 100 m. Temperatures were recorded in putative microrefugia and in neighboring plots for approximately 2 years to quantify their thermal offsets. Vascular plant inventories were carried out to test whether plant communities also reflect microclimatic offsets. We found that current microclimatic dynamics are genuinely at stake in microrefugia. Microrefugia climates are systematically colder compared to those found in neighboring control plots. This pattern was more noticeable during the summer months. Abyssal populations showed stronger offsets compared to neighboring plots than the putative microrefugia occurring at higher altitudes. Plant communities demonstrate this strong spatial climatic variability, even at such a microscale approach, as species compositions systematically differed between the two plots, with species more adapted to colder and moister conditions in microrefugia compared to the surrounding area.

Coronilla juncea, a native candidate for phytostabilization of potentially toxic elements and restoration of Mediterranean soils.

Soil contamination pattern due to industrial activities often leads to high concentrations of potentially toxic elements (PTE) decreasing with depth. This spatial heterogeneity of the soil contamination may have significant consequences on the soil properties and soil living communities. We evaluated the effects of both surface and solum soil contamination heterogeneity on Coronilla juncea L. (Fabaceae) functional traits in field conditions and the phytostabilization potential of this species. Plant and soil samples were collected on 3 sites along a PTE contamination gradient. The correlations between PTE concentration in plant and soil samples at 2 depths, physico-chemical properties of soil, plant biomass and soil microbial activity were tested. Field measurements highlight a decreasing PTE concentration with soil depth in addition to an important surface heterogeneity of As, Cu, Pb, Sb and Zn soil concentrations. Root PTE concentrations in C. juncea did not follow soil PTE concentrations. Concentrations of PTE in the root parts were higher than those of the aerial parts. Low PTE translocation and root symbioses with microorganisms suggest that this native plant species may play a role as engineer species with positive implications for the phytostabilization of Mediterranean PTE contaminated soils and their ecological restoration.

Climate shapes the seed germination niche of temperate flowering plants: a meta-analysis of European seed conservation data.

BACKGROUND AND AIMS: Interactions between ecological factors and seed physiological responses during the establishment phase shape the distribution of plants. Yet, our understanding of the functions and evolution of early-life traits has been limited by the scarcity of large-scale datasets. Here, we tested the hypothesis that the germination niche of temperate plants is shaped by their climatic requirements and phylogenetic relatedness, using germination data sourced from a comprehensive seed conservation database of the European flora (ENSCOBASE). METHODS: We performed a phylogenetically informed Bayesian meta-analysis of primary data, considering 18 762 germination tests of 2418 species from laboratory experiments conducted across all European geographical regions. We tested for the interaction between species' climatic requirements and germination responses to experimental conditions including temperature, alternating temperature, light and dormancy-breaking treatments, while accounting for between-study variation related to seed sources and seed lot physiological status. KEY RESULTS: Climate was a strong predictor of germination responses. In warm and seasonally dry climates the seed germination niche includes a cold-cued germination response and an inhibition determined by alternating temperature regimes and cold stratification, while in climates with high temperature seasonality opposite responses can be observed. Germination responses to scarification and light were related to seed mass but not to climate. We also found a significant phylogenetic signal in the response of seeds to experimental conditions, providing evidence that the germination niche is phylogenetically constrained. Nevertheless, phylogenetically distant lineages exhibited common germination responses under similar climates. CONCLUSION: This is the first quantitative meta-analysis of the germination niche at a continental scale. Our findings showed that the germination niches of European plants exhibit evolutionary convergence mediated by strong pressures at the macroclimatic level. In addition, our methodological approach highlighted how large datasets generated by conservation seed banking can be valuable sources to address questions in plant macroecology and evolution.

Amplified drought induced by climate change reduces seedling emergence and increases seedling mortality for two Mediterranean perennial herbs.

Seedling recruitment is a bottleneck for population dynamics and range shift. The vital rates linked to recruitment by seed are impacted by amplified drought induced by climate change. In the Mediterranean region, autumn and winter seedling emergence and mortality may have strong impact on the overall seedling recruitment. However, studies focusing on the temporal dynamic of recruitment during these seasons are rare. This study was performed in a deciduous Mediterranean oak forest located in southern France and quantifies the impact of amplified drought conditions on autumn and winter seedling emergence and seedling mortality rates of two herbaceous plant species with meso-Mediterranean and supra-Mediterranean distribution (respectively, Silene italica and Silene nutans). Seedlings were followed from October 2019 to May 2020 in both undisturbed and disturbed plots where the litter and the aboveground biomass have been removed to create open microsites. Amplified drought conditions reduced seedling emergence and increased seedling mortality for both Silene species but these negative effects were dependent on soil disturbance conditions. Emergence of S. italica decreased only in undisturbed plots (-7%) whereas emergence of S. nutans decreased only in disturbed plots (-10%) under amplified drought conditions. The seedling mortality rate of S. italica was 51% higher under amplified drought conditions in undisturbed plots while that of S. nutans was 38% higher in disturbed plots. Aridification due to lower precipitation in the Mediterranean region will negatively impact the seedling recruitment of these two Silene species. Climate change effects on early vital rates may likely have major negative impacts on the overall population dynamic.

The seed germination spectrum of alpine plants: a global meta-analysis.

Assumptions about the germination ecology of alpine plants are presently based on individual species and local studies. A current challenge is to synthesise, at the global level, the alpine seed ecological spectrum. We performed a meta-analysis of primary data from laboratory experiments conducted across four continents (excluding the tropics) and 661 species, to estimate the influence of six environmental cues on germination proportion, mean germination time and germination synchrony; accounting for seed morphology (mass, embryo : seed ratio) and phylogeny. Most alpine plants show physiological seed dormancy, a strong need for cold stratification, warm-cued germination and positive germination responses to light and alternating temperatures. Species restricted to the alpine belt have a higher preference for warm temperatures and a stronger response to cold stratification than species whose distribution extends also below the treeline. Seed mass, embryo size and phylogeny have strong constraining effects on germination responses to the environment. Globally, overwintering and warm temperatures are key drivers of germination in alpine habitats. The interplay between germination physiology and seed morphological traits further reflects pressures to avoid frost or drought stress. Our results indicate the convergence, at the global level, of the seed germination patterns of alpine species.

A research agenda for seed-trait functional ecology.

Trait-based approaches have improved our understanding of plant evolution, community assembly and ecosystem functioning. A major challenge for the upcoming decades is to understand the functions and evolution of early life-history traits, across levels of organization and ecological strategies. Although a variety of seed traits are critical for dispersal, persistence, germination timing and seedling establishment, only seed mass has been considered systematically. Here we suggest broadening the range of morphological, physiological and biochemical seed traits to add new understanding on plant niches, population dynamics and community assembly. The diversity of seed traits and functions provides an important challenge that will require international collaboration in three areas of research. First, we present a conceptual framework for a seed ecological spectrum that builds upon current understanding of plant niches. We then lay the foundation for a seed-trait functional network, the establishment of which will underpin and facilitate trait-based inferences. Finally, we anticipate novel insights and challenges associated with incorporating diverse seed traits into predictive evolutionary ecology, community ecology and applied ecology. If the community invests in standardized seed-trait collection and the implementation of rigorous databases, major strides can be made at this exciting frontier of functional ecology.

Islands, mainland, and terrestrial fragments: How isolation shapes plant diversity.

The fragmentation of natural habitats is a major threat for biodiversity. However, the impact and spatial scale of natural isolation mechanisms leading to species loss, compared to anthropogenic fragmentation, are not clear, mainly due to differences between fragments and islands, such as matrix permeability. We studied a 500 km2 Mediterranean region in France, including urban habitat fragments, continuous habitat, and continental-shelf islands. On the basis of 295 floristic relevés, we built species-area relationships to compare isolation in fragments after urbanization, with continuous habitat and continental-shelf islands. We assumed either no dispersal, infinite dispersal, or estimated intermediate levels of habitat reachability through graph theory. Isolation mechanisms occurred in fragments but with a lower strength than in near-shore islands, and most importantly affected perennial plants. Annual plants were less affected, probably due to their smaller size and shorter life cycle. Isolation occurred at landscape level in fragments and at patch level in islands. The amount of reachable habitat (accounting for spatial configuration) explained local species richness in both systems, but the amount of habitat (no consideration of spatial configuration) was already a good predictor. These results suggest an important role of habitat amount around fragments in mitigating the isolation effects observed in near-shore islands, and the importance of carefully considering different functional groups.

Prioritizing conservation areas for coastal plant diversity under increasing urbanization.

Coastal urban expansion will continue to drive further biodiversity losses, if conservation targets for coastal ecosystems are not defined and met. Prioritizing areas for future protected area networks is thus an urgent task in such urbanization-threatened ecosystems. Our aim is to quantify past and future losses of coastal vegetation priority areas due to urbanization and assess the effectiveness of the existing protected area network for conservation. We conduct a prioritization analysis, based on 82 coastal plants, including common and IUCN red list species, in a highly-urbanized but biotically diverse region, in South-Eastern France. We evaluate the role of protected areas, by taking into account both strict and multi-use areas. We assess the impact of past and future urbanization on high priority areas, by combining prioritization analyses and urbanization models. We show that half of the highly diverse areas have already been lost due to urbanization. Remaining top priority areas are also among the most exposed to future urban expansion. The effectiveness of the existing protected area (PA) network is only partial. While strict PAs coincide well with top priority areas, they only represent less than one third of priority areas. The effectiveness of multi-use PAs, such as the Natura 2000 network, also remains limited. Our approach highlights the impact of urbanization on plant conservation targets. By modelling urbanization, we manage to identify those areas where protection could be more efficient to limit further losses. We suggest to use our approach in the future to expand the PA network in order to achieve the 2020 Aichi biodiversity targets.

Identification and prioritization of areas with high environmental risk in Mediterranean coastal areas: A flexible approach.

Interdisciplinarity and transdisciplinarity are the cornerstone for the future management of coastal ecosystems with many vulnerability and hazard indexes developed for this purpose, especially in the engineering literature, but with limited studies that considered ecological implications within a risk assessment. Similarly, the concept of prioritization of sites has been widely examined in biodiversity conservation studies, but only recently as an instrument for territory management. Considering coastal plant diversity at the species and community levels, and their vulnerability to three main potential hazards threatening coastal areas (oil spills, Hazardous and Noxious Substances pollution, fragmentation of natural habitats), the objective of this paper is to define an easy-to-use approach to locate and prioritize the areas more susceptible to those stressors, in order to have a practical instrument for risk management in the ordinary and extra-ordinary management of the coastline. The procedure has been applied at pilot areas in four Mediterranean countries (Italy, France, Lebanon and Tunisia). This approach can provide policy planners, decision makers and local communities an easy-to-use instrument able to facilitate the implementation of the ICZM (Integrated Coastal Zone Management) process in their territory.

Germination traits explain soil seed persistence across species: the case of Mediterranean annual plants in cereal fields.

BACKGROUND AND AIMS: Seed persistence in the soil under field conditions is an important issue for the maintenance of local plant populations and the restoration of plant communities, increasingly so in the light of rapidly changing land use and climate change. Whereas processes important for dispersal in space are well known, knowledge of processes governing dispersal in time is still limited. Data for morphological seed traits such as size have given contradictory results for prediction of soil seed persistence or cover only a few species. There have been few experimental studies on the role of germination traits in determining soil seed persistence, while none has studied their predictive value consistently across species. Delayed germination, as well as light requirements for germination, have been suggested to contribute to the formation of persistent seed banks. Moreover, diurnally fluctuating temperatures can influence the timing of germination and are therefore linked to seed bank persistence. METHODS: The role of germination speed measured by T(50) (days to germination of 50 % of all germinated seeds), light requirement and reaction to diurnally fluctuating temperatures in determining seed persistence in the soil was evaluated using an experimental comparative data set of 25 annual cereal weed species. KEY RESULTS: It is shown that light requirements and slow germination are important features to maintain seeds ungerminated just after entering the soil, and hence influence survival of seeds in the soil. However, the detection of low diurnally fluctuating temperatures enhances soil seed bank persistence by limiting germination. Our data further suggest that the effect of diurnally fluctuating temperatures, as measured on seeds after dispersal and dry storage, is increasingly important to prevent fatal germination after longer burial periods. CONCLUSIONS: These results underline the functional role of delayed germination and light for survival of seeds in the soil and hence their importance for shaping the first part of the seed decay curve. Our analyses highlight the detection of diurnally fluctuating temperatures as a third mechanism to achieve higher soil seed persistence after burial which interacts strongly with season. We therefore advocate focusing future research on mechanisms that favour soil seed persistence after longer burial times and moving from studies of morphological features to exploration of germination traits such as reaction to diurnally fluctuating temperatures.

The seed bank longevity index revisited: limited reliability evident from a burial experiment and database analyses.

BACKGROUND AND AIMS: Seed survival in the soil contributes to population persistence and community diversity, creating a need for reliable measures of soil seed bank persistence. Several methods estimate soil seed bank persistence, most of which count seedlings emerging from soil samples. Seasonality, depth distribution and presence (or absence) in vegetation are then used to classify a species' soil seed bank into persistent or transient, often synthesized into a longevity index. This study aims to determine if counts of seedlings from soil samples yield reliable seed bank persistence estimates and if this is correlated to seed production. METHODS: Seeds of 38 annual weeds taken from arable fields were buried in the field and their viability tested by germination and tetrazolium tests at 6 month intervals for 2.5 years. This direct measure of soil seed survival was compared with indirect estimates from the literature, which use seedling emergence from soil samples to determine seed bank persistence. Published databases were used to explore the generality of the influence of reproductive capacity on seed bank persistence estimates from seedling emergence data. KEY RESULTS: There was no relationship between a species' soil seed survival in the burial experiment and its seed bank persistence estimate from published data using seedling emergence from soil samples. The analysis of complementary data from published databases revealed that while seed bank persistence estimates based on seedling emergence from soil samples are generally correlated with seed production, estimates of seed banks from burial experiments are not. CONCLUSIONS: The results can be explained in terms of the seed size-seed number trade-off, which suggests that the higher number of smaller seeds is compensated after germination. Soil seed bank persistence estimates correlated to seed production are therefore not useful for studies on population persistence or community diversity. Confusion of soil seed survival and seed production can be avoided by separate use of soil seed abundance and experimental soil seed survival.