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A knowledge of the morphology of seeds and diaspores is an invaluable resource that has prompted the compilation of descriptions and/or images in a variety of media, including databases with standardized protocols for trait measurement. The pressing need for new solutions to solve environmental crises, together with advances in the application of machine learning and image analysis to solve ecological problems, call for an upgrade of the diaspore morphological datasets. Here, we present DiasMorph, a dataset of morphological traits and images of diaspores from 1,437 taxa in 513 genera, and 96 families from Central Europe, totalling 94,214 records. Moreover, DiasMorph compiles information on quantitative morphological traits extracted from the images following a standardized and reproducible image analysis method, and include not only traditional morphometric measurements, but also colour, and shape features made available for the first time in a large dataset. The quantitative traits records correspond to measurements of individual diaspores, an input currently unavailable in traits databases that will allow for several approaches to be used for a complete exploration of the morphological traits of these species. We also included information on the presence and absence of appendages and structures in the diaspores of the evaluated taxa. By making these data available, we aim to encourage initiatives to advance on new tools for diaspore identification, further our understanding of morphological traits functions, and provide means for the continuous development of image analyses applications. Keywords: diaspore appendages, dispersal unit, image dataset, plant traits dataset, seed colour, seed morphology, seed shape, seed size, seed traits.

Reassembling ecological communities and rebuilding habitats through active restoration treatments require curating the selection of plant species to use in seeding and planting mixes. Ideally, these mixes should be assembled based on attributes that support ecosystem function and services, promote plant and animal species interactions and ecological networks in restoration while balancing project constraints. Despite these critical considerations, it is common for species mixes to be selected opportunistically. Reframing the selection of seed mixes for restoration around ecological objectives is essential for success but accessible methods and tools are needed to support this effort. We developed a framework to optimize species seed mixes based on prioritizing plant species attributes to best support different objectives for ecosystem functions, services and trophic relationships such as pollination, seed dispersal and herbivory. We compared results to approaches where plant species are selected to represent plant taxonomic richness, dominant species and at random. We tested our framework in European alpine grasslands by identifying 176 plant species characteristic of the species pool, and identified 163 associated attributes affiliated to trophic relationships, ecosystem functions and services. In all cases, trophic relationships, ecosystem functions and services can be captured more efficiently through objective-based prioritization using the functional identity of plant species. Solutions (plant species lists) can be compared quantitatively, in terms of costs, species or objectives. We confirm that a random draw of plant species from the regional plant species pool cannot be assumed to support other trophic groups and ecosystem functions and services. Synthesis and Applications. Our framework is presented as a proof-of-concept to help restoration practitioners better apply quantitative decision support to plant species selection to efficiently meet ecological restoration outcomes. Our approach may be tailored to any restoration initiative, habitat or restoration targets where seeding or planting mixes will be applied in active treatments. As global priority and resources are increasingly placed into restoration, this approach could be advanced to help make efficient decisions for many stages of the restoration process.

Species may benefit from green infrastructure, i.e. the network of natural and anthropogenic habitat remnants in human-dominated landscapes, if it helps isolated populations in remaining habitat patches to be functionally connected. The importance of green infrastructure is therefore increasingly emphasized in conservation policy to counter biodiversity loss. However, there is limited evidence, particularly in plants, that green infrastructure promotes functional connectivity, i.e. supports the colonization of habitat patches across a landscape. We applied landscape genetics to test whether the green infrastructure supports structural and functional connectivity in the grassland perennial Galium verum, in 35 landscapes in Belgium, Germany and Sweden. We used multivariate genetic clustering techniques, nestedness analyses and conditional inference trees to examine landscape-scale patterns in genetic diversity and structure of plant populations in the green infrastructure surrounding semi-natural grasslands. Inferred functional connectivity explained genetic variation better than structural connectivity, yielding positive effects on genetic variation. The road verge network, a major structural component of the green infrastructure and its functional connectivity, most effectively explained genetic diversity and composition in G. verum. Galium verum ramets occupying the surrounding landscape proved to be genetic subsets of focal grassland populations, shaping a nested landscape population genetic structure with focal grasslands, particularly ancient ones, harbouring unique genetic diversity. This nested pattern weakened as road network density increased, suggesting road verge networks enable high landscape occupancy by increased habitat availability and facilitates gene flow into the surrounding landscape. Our study proposes that green infrastructure can promote functional connectivity, providing that a plant species can survive outside of core habitat patches. As this often excludes habitat specialist species, conservation practice and policy should primarily focus on ancient, managed semi-natural grasslands. These grasslands both harbour unique genetic diversity and act as primary gene and propagule sources for the surrounding landscape, highlighting their conservation value., This research was funded through the 2015–2016 BiodivERsA COFUND call for research proposals, with the national funders FORMAS, the Swedish Environmental Protection Agency (Naturvårdsverket), the Belgian Science Policy Office (BelSPo), the Germany Federal Ministry of Education and Research (Bundesministerium fuer Bildung und Forschung, FKZ: 01LC1619A) and the Spanish Ministry of Science, Innovation and Universities (Ministerio de Ciencia, Innovación y Universidades). JMB was funded under UKCEH National Capability project no. 06895.

Long-term analyses of biodiversity data highlight a 'biodiversity conservation paradox': biological communities show substantial species turnover over the past century1,2, but changes in species richness are marginal1,3-5. Most studies, however, have focused only on the incidence of species, and have not considered changes in local abundance. Here we asked whether analysing changes in the cover of plant species could reveal previously unrecognized patterns of biodiversity change and provide insights into the underlying mechanisms. We compiled and analysed a dataset of 7,738 permanent and semi-permanent vegetation plots from Germany that were surveyed between 2 and 54 times from 1927 to 2020, in total comprising 1,794 species of vascular plants. We found that decrements in cover, averaged across all species and plots, occurred more often than increments; that the number of species that decreased in cover was higher than the number of species that increased; and that decrements were more equally distributed among losers than were gains among winners. Null model simulations confirmed that these trends do not emerge by chance, but are the consequence of species-specific negative effects of environmental changes. In the long run, these trends might result in substantial losses of species at both local and regional scales. Summarizing the changes by decade shows that the inequality in the mean change in species cover of losers and winners diverged as early as the 1960s. We conclude that changes in species cover in communities represent an important but understudied dimension of biodiversity change that should more routinely be considered in time-series analyses.

The conservation of genetic diversity requires a deeper understanding of the processes shaping intraspecific variation. In recent decades, numerous studies identified various drivers of genetic diversity. However, the interpretation of these drivers remains inconsistent, since their strength and balance may differ between species and habitats. Therefore, we analyzed potential drivers of genetic diversity in a comparative multispecies approach across different semi-natural grassland types. We used molecular markers to detect genetic diversity in populations of nine typical grassland plant species from dry calcareous grasslands, mesic hay meadows, and wet litter meadows in Central Europe. Additionally, we collected data on potential drivers of genetic diversity, which were assigned to four categories describing habitat age, landscape structure, habitat quality, and population size. Subsequently, we applied multiple linear regression models and variation partitioning analyses to identify the most influential drivers of genetic diversity in semi-natural grassland plant populations. Our study revealed clear differences in drivers of genetic diversity between grassland types. In calcareous grasslands genetic diversity depended almost completely on landscape structure. However, we identified habitat age and habitat quality as additional drivers in hay meadows, while population size was another driver in litter meadows. The strong variation in drivers of genetic diversity in hay and litter meadows can be ascribed to higher levels of environmental variation among these sites and due to their more recent origin than calcareous grassland sites. We conclude that different drivers of genetic diversity must be considered to maintain high levels of intraspecific diversity in different grassland types.

Millenia of sustainable, low intensity land use have formed the cultural landscapes of central Europe. Studies from the Central Alps show that mountain pastures also look back onto many thousand years of land use history. In this palynological and pedoanthracological study in the border region between Germany and Austria in the Mangfall Mountains, we aim to close the knowledge gap that exists for the German part of the Northern Alps, where no conclusive evidence for the onset of pastoral activities has been presented so far. Our results reveal strong evidence, that mountain pasture use in this region reaches back to the Iron Age at least. However, the reconstruction of vegetation and fire history indicates human interaction with the environment much earlier, starting in the Neolithic Age, where we found evidence of slash and burn activities and first occurrences of pasture indicator pollen. A rising number of mega charcoal pieces dated to the Bronze Age suggests increased slash and burn activities, possibly linked to the creation of open space for pasturing. Therefore, our results provide profound evidence of human interaction with the mountain environment, beginning in the Neolithic Age and clear evidence of mountain pasture use beginning during the Iron Age at 750 BC. Based on palynology and pedoanthracology it is, however, difficult to clearly differentiate between pasturing, hunting and other human interactions with the environment. Further archaeological studies in this area could add valuable information to our findings and shed more light onto the early history of farming activities in the Northern Alps.

[Context] Functional connectivity is vital for plant species dispersal, but little is known about how habitat loss and the presence of green infrastructure interact to affect both functional and structural connectivity, and the impacts of each on species groups., [Objectives] We investigate how changes in the spatial configuration of species-rich grasslands and related green infrastructure such as road verges, hedgerows and forest borders in three European countries have influenced landscape connectivity, and the effects on grassland plant biodiversity., [Methods] We mapped past and present land use for 36 landscapes in Belgium, Germany and Sweden, to estimate connectivity based on simple habitat spatial configuration (structural connectivity) and accounting for effective dispersal and establishment (functional connectivity) around focal grasslands. We used the resulting measures of landscape change to interpret patterns in plant communities., [Results] Increased presence of landscape connecting elements could not compensate for large scale losses of grassland area resulting in substantial declines in structural and functional connectivity. Generalist species were negatively affected by connectivity, and responded most strongly to structural connectivity, while functional connectivity determined the occurrence of grassland specialists in focal grasslands. Restored patches had more generalist species, and a lower density of grassland specialist species than ancient patches., [Conclusions] Protecting both species rich grasslands and dispersal pathways within landscapes is essential for maintaining grassland biodiversity. Our results show that increases in green infrastructure have not been sufficient to offset loss of semi-natural habitat, and that landscape links must be functionally effective in order to contribute to grassland diversity., This research was funded through the 2015–2016 BiodivERsA COFUND call for research proposals, with the national funders the Swedish Research Council for Environment, Agricultural Sciences and Spatial Planning (FORMAS), the Swedish Environmental Protection Agency (Naturvårdsverket), the Belgian Science Policy Office (BelSPo), the Germany Federal Ministry of Education and Research (Bundesministerium fuer Bildung und Forschung, FKZ: 01LC1619A) and the Spanish Ministry of Science, Innovation and Universities (Ministerio de Ciencia, Innovación y Universidades). Open access funding provided by Stockholm University.

Mudflats are exposed for short periods after flood water drawdown. They support fast-growing annual herbs with a ruderal strategy. To optimize their recruitment success, seeds of mudflat species germinate better under fluctuating temperatures, full illumination and aerobic environments that indicate the presence of optimal (non-flooded) conditions for plant growth and development. Here, we hypothesize that prior exposure of mudflat seeds to hypoxic (flooded) environment interferes with the germination process and results in more vigorous germination once aerobic conditions are regained. To test this hypothesis, seeds of five mudflat species were incubated in both aerobic and hypoxic environments at four (14/6, 22/14, 22/22 and 30/22°C) temperature regimes, reflecting different (seasonal) conditions when drawdowns may occur. All species responded positively to four temperature regimes; however, moderate 22/14 and 22/22°C temperatures were optimum for high percentages and rates (speed) of seed germination. Since seeds of four species germinated exclusively under aerobic conditions, they were moved from hypoxic to aerobic conditions. Prior exposure of seeds to hypoxic environment facilitated high percentages, rates and synchronization of germination of Limosella aquatica, Peplis portula and Samolus valerandi seeds compared to incubation under strict aerobic conditions. However, prior exposure to hypoxic environment induced secondary dormancy in non-dormant seeds of Hypericum humifusum but broke dormancy in Lythrum hyssopifolia seeds that otherwise required cold stratification to overcome physiological dormancy. All species that have a narrow ecological niche (strictly occurring in mudflat habitats) showed positive responses to prior exposure to hypoxic environments. In contrast, H. humifusum that has a wide ecological niche (from mudflats to moist sandy grasslands) showed a negative response. We conclude that the hypoxic environment may strongly affect seed germination behaviour once the aerobic environment is regained. The most striking effect is the acceleration of the germination process and, therefore, life cycle supporting the survival in an ephemeral habitat.

In a continuous, perfectly stratified sedimentary sequence which was discovered under a large sandstone overhang in northern Bohemia, Czech Republic, we analysed multiple biological remains, archaeological features and artefacts. This multi-proxy record has allowed us to examine the interactions between woodland and humans in a permanently wooded environment throughout almost the entire Holocene. We paid most attention to massive finds of dung pellets from sheep, goats or pigs and bedding layers which show that the site was used as a pen and shelter for livestock which grazed in the woods. Our results imply that such practices have occurred since the Neolithic, but the most robust evidence of these is for the Iron Age and early Middle Ages. Detailed analyses of the dung indicate woodland grazing and foddering with branches, acorns, beechnuts and crop processing remains. In addition, the wide palaeoenvironmental range of this detailed investigation provides evidence of the impact of wood pasturing on ecological functions, taxon composition and diversity of the local woodland ecosystem in the Holocene.

Information on the regeneration ecology of native woody species of the forest-savanna ecotone of West Africa is scarce, which is a major impediment to their optimal utilization in large-scale restoration programmes. The scattered information that is available for some of these species reveals that freshly matured seed are dormant. However, environmental heterogeneity among different habitats may results in inter-population seed dormancy variation. Thus, our objective was to re-examine the dormancy of 12 species from the forest-savanna ecotone that have been targeted for reforestation. Specifically, we aimed to examine the water-permeability of the seeds and explore the effectiveness of acid scarification and heat treatment to alleviate dormancy. Four species belonging to families other than Fabaceae and Malvaceae had water-permeable seeds. Two of them had nondormant (ND) seeds, and seeds of the other two species had a mixture of ND and other kinds of dormancy (possibly physiological dormancy, PD). Most species of Fabaceae and Malvaceae had water-impermeable seeds. All seeds of three species had physical dormant (PY), and some seeds of the remaining species had PY, while others were ND or had PD. Acid-scarification was effective in breaking PY and in augmenting imbibition and germination of non-PY seeds, while heat treatment was moderately effective in breaking dormancy. In general, acid scarification for 1-30 minutes and heat treatment for one hour at 55-75°C were optimal to enhance seed germination, depending on species. The present study has wide practical implications for park conservationists and restoration ecologists interested in producing bulk quantities of high-quality planting stocks of native woody species for large-scale restoration programmes.

QuestionSoil seed banks (SSB), i.e., pools of viable seeds in the soil and its surface, play a crucial role in plant biology and ecology. Information on seed persistence in soil is of great importance for fundamental and applied research, yet compiling datasets on this trait still requires enormous efforts. We asked whether the machine learning (ML) approach could be used to infer and predict SSB properties of a regional flora based on easily available data.LocationEighteen calcareous grasslands located along an elevational gradient of almost 2000 m in the Bavarian Alps, Germany.MethodsWe compared a commonly used ML model (random forest) with a conventional model (linear regression model) as to their ability to predict SSB presence/absence and density using empirical data on SSB characteristics (environmental, seed traits and phylogenetic predictors). Further, we identified the most important determinants of seed persistence in soil for predicting qualitative and quantitative SSB characteristics using the ML approach.ResultsWe demonstrated that the ML model predicts SSB characteristics significantly better than the linear regression model. A single set of predictors (either environment, or seed traits, or phylogenetic eigenvectors) was sufficient for the ML model to achieve high performance in predicting SSB characteristics. Importantly, we established that a few widely available SSB predictors can achieve high predictive power in the ML approach, suggesting a high flexibility of the developed approach for use in various study systems.ConclusionsOur study provides a novel methodological approach that combines empirical knowledge on the determinants of SSB characteristics with a modern, flexible statistical approach based on ML. It clearly demonstrates that ML can be developed into a key tool to facilitate labor-intensive, costly and time-consuming functional trait research.

Assembly of plant communities has long been scrutinized through the lens of trait-based ecology. Studies generally analyze functional traits related to the vegetative growth, survival and resource acquisition and thus ignore how ecological processes may affect plants at other stages of their lifecycle, particularly when seeds disperse, persist in soil and germinate. Here, we analyzed an extensive data set of 16 traits for 167 species measured in-situ in 36 grasslands located along an elevational gradient and compared the impact of abiotic filtering, biotic interactions and dispersal on traits reflecting different trait categories: plant vegetative growth, germination, dispersal, and seed morphology. For each community, we quantified community weighted mean (CWM) and functional diversity (FD) for all traits and established their relationships to mean annual temperature. The seed traits were weakly correlated to vegetative traits. Therefore, these traits constituted independent axes of plant phenotypical variation affected differently by the ecological processes considered. Abiotic filtering impacted mostly the vegetative traits and to a lesser extent on seed germination and morphological traits. Increasing low-temperature stress towards colder sites selected for short-stature, slow-growing and frost-tolerant species that produce small quantity of smaller seeds with higher degree of dormancy, high temperature requirements for germination and comparatively low germination speed. Biotic interactions, specifically competition in the lowlands and facilitation in uplands, also filtered certain functional traits in the study communities. The benign climate in lowlands promoted plant with competitive strategies including fast growth and resource acquisition (vegetative growth traits) and early and fast germination (germination traits), whereas the effects of facilitation on the vegetative and germination traits were cancelled out by the strong abiotic filtering. The changes in the main dispersal vector from zoochory to anemochory along the gradient strongly affected the dispersal and the seed morphological trait structure of the communities. Specifically, stronger vertical turbulence and moderate warm-upwinds combined with low grazing intensity selected for light and non-round shaped seeds with lower terminal velocity and endozoochorous potential. Synthesis. We clearly demonstrate that, in addition to vegetative traits, seed traits can substantially contribute to functional structuring of plant communities along environmental gradients. Thus, the ‚hard‘ seed traits related to germination and dispersal are critical to detect multiple, complex community assembly rules. Consequently, such traits should be included in core lists of plant traits and, when applicable, be incorporated into analysis of community assembly. Assembly of plant communities has long been scrutinized through the lens of trait-based ecology. Studies generally analyze functional traits related to the vegetative growth, survival and resource acquisition and thus ignore how ecological processes may affect plants at other stages of their lifecycle, particularly when seeds disperse, persist in soil and germinate. Here, we analyzed an extensive data set of 16 traits for 167 species measured in-situ in 36 grasslands located along an elevational gradient and compared the impact of abiotic filtering, biotic interactions and dispersal on traits reflecting different trait categories: plant vegetative growth, germination, dispersal, and seed morphology. For each community, we quantified community weighted mean (CWM) and functional diversity (FD) for all traits and established their relationships to mean annual temperature. The seed traits were weakly correlated to vegetative traits. Therefore, these traits constituted independent axes of plant phenotypical variation affected differently by the ecological processes considered. Abiotic filtering impacted mostly the vegetative traits and to a lesser extent on seed germination and morphological traits. Increasing low-temperature stress towards colder sites selected for short-stature, slow-growing and frost-tolerant species that produce small quantity of smaller seeds with higher degree of dormancy, high temperature requirements for germination and comparatively low germination speed. Biotic interactions, specifically competition in the lowlands and facilitation in uplands, also filtered certain functional traits in the study communities. The benign climate in lowlands promoted plant with competitive strategies including fast growth and resource acquisition (vegetative growth traits) and early and fast germination (germination traits), whereas the effects of facilitation on the vegetative and germination traits were cancelled out by the strong abiotic filtering. The changes in the main dispersal vector from zoochory to anemochory along the gradient strongly affected the dispersal and the seed morphological trait structure of the communities. Specifically, stronger vertical turbulence and moderate warm-upwinds combined with low grazing intensity selected for light and non-round shaped seeds with lower terminal velocity and endozoochorous potential. Synthesis. We clearly demonstrate that, in addition to vegetative traits, seed traits can substantially contribute to functional structuring of plant communities along environmental gradients. Thus, the ‚hard‘ seed traits related to germination and dispersal are critical to detect multiple, complex community assembly rules. Consequently, such traits should be included in core lists of plant traits and, when applicable, be incorporated into analysis of community assembly.

Ancient semi-natural grasslands in Europe are important for ecosystem service (ES) provision. Often, the surrounding matrix contains 'Grassland Green Infrastructure' (GGI) that contain grassland species which have the potential to supplement grassland ES provision across the landscape. Here we investigate the potential for GGI to deliver a set of complementary ES, driven by plant composition.We surveyed 36 landscapes across three European countries comprising core grasslands and their surrounding GGI. We calculated community-level values of plant species characteristics to provide indicators for four ES: nature conservation value, pollination, carbon storage and aesthetic appeal.Inferred ES delivery for GGI was substantially lower than in core grasslands for conservation, pollination and aesthetic appeal indicators, but not for carbon storage. These differences were driven by the GGI having 17% fewer plant species, and compositional differences, with 61% of species unique to the core grasslands. In addition, connectivity to the core, the amount of GGI and inferred seed dispersal distances by livestock, were strongly positively correlated with conservation value, pollination and aesthetic indicators. All ES indicators showed similar responses to the GGI spatial structure and distance to the core, suggesting robust effects of these drivers on ES. We projected that improved landscape-wide delivery of nature conservation value and pollination could be achieved through targeted GGI management. Reductions in the distances seeds would need to disperse, more GGI, along with a diversification of the GGI elements, were predicted to enhance service credits.We conclude that for vegetation-related ES, species surveys can be employed to assess potential ES delivery. Creating and enhancing GGI is a useful landscape management strategy to supplement the ES delivered by ancient grasslands.

Central European grasslands, such as calcareous grasslands and oat-grass meadows, are characterized by diverse environmental conditions and management regimes. Therefore, we aimed to determine potential differences in genetic and epigenetic variation patterns between the contrasting habitats and to identify the drivers of genetic and epigenetic variation. We investigated the genetic and epigenetic variation of the ecologically variable plant species

Species from OCBILs (old, climatically buffered, infertile landscapes) are expected to exhibit reduced dispersability, linked to long-term stable environmental conditions. We tested the prediction of limited dispersability in plant communities in campo rupestre, a megadiverse OCBIL grassland. In a field experiment, we sampled sticky and funnel traps for 12 months along paired preserved and disturbed plots to estimate and compare annual seed density and richness. Our community-level seed rain assessment suggested widespread seed limitation in both preserved and disturbed areas, expressed by the low seed density in traps (average of 2.6 seeds/m2/day) and remarkably low seed density found for 90% of the morphospecies. Although seed density in campo rupestre was much lower than for other grasslands, species richness (92 seed morphospecies) was the highest reported in the literature. Higher seed density and species richness were captured in funnel traps, especially during the rainy season. Approximately 32% of morphospecies were registered exclusively in disturbed plots and 25% exclusively in preserved plots. This limited seed dispersability is of particular relevance for a better understanding of seed ecology and resilience in campo rupestre and may partially explain the poor recovery of these disturbed areas after soil removal.

Dry calcareous grasslands are among the most species-rich habitats in Central Europe, harbouring numerous threatened species. Because of their strong decline, they are being protected under the European Habitats Directive. However, apart from this general decline, new grasslands developed after the abandonment of arable fields on marginal land over the course of the last few centuries or even decades. The main question of this study was which species may indicate the age of a dry calcareous grassland habitat in the Franconian Jurassic mountains near Kallmunz. We compared the diversity parameters and nature conservation value of both grassland types. Additionally, we searched for differences in habitat and soil parameters. Furthermore, we asked if there is a general pattern of indicator species among available studies on ancient (i.e. those continuously used as pastures at least since 1830) and recent (i.e. those temporarily farmed as arable fields after 1830) calcareous grasslands. We compiled a list of indicator species of both ancient and recent grasslands in the study region. Comparison with other studies leads to the conclusion that there are not many species that clearly indicate grassland age across different regions (the best indicators are Carex caryophyllea, Cirsium acaule and Hippocrepis comosa for ancient grasslands, and Agrimonia eupatoria and Astragalus glyciplyllos for recent grasslands). Ancient grasslands harbour a somewhat greater number of threatened species than recent grasslands. Many species of the ancient grasslands under study can be considered relict species of steppic grasslands or open pine forests (e.g. Hippocrepis comosa, Pulsatilla vulgaris, Teucrium chamaedrys, Teucrium montanum and Thymus praecox). Recent grasslands also harbour rare and endangered species, especially disturbance-tolerant relicts of former arable use (e.g. Melampyrum arvense) and may therefore be of high conservation value, too. The average number of species per plot is greater in ancient grasslands. However, the most species-rich plot (46 species of vascular plants within a 4-m2 quadrat) was found in a 60 years old grassland. Arable cultivation in the past has altered the physical and chemical properties of the soil of recent grasslands. In general, ancient grasslands occur on nutrient-poorer and less calcium-rich soils with high water holding capacity. High water holding capacity is connected with high humus content, which increases the importance of ancient grasslands for carbon storage. The challenges and benefits of differentiating grasslands of different age in the management of protected areas and landscape planning (e.g. the identification of High Nature Value farmland) are discussed.

Alpine farming and pasturing at high altitudes in the Alps has created one of the most species-rich and diverse landscapes in Europe. In order to fully understand, appreciate and protect these habitats it is essential to learn about their history and origin. Until the present day, alpine farming provides essential additional food sources for livestock of farmers in the alpine valleys. Based on written sources, historians are able to track alpine farming back to the Middle Ages. Other approaches from different fields in science, however, can look back even further in search of evidence for alpine farming. This interdisciplinary literature review therefore aims to summarize the scientific work that has been done in different fields of science such as Archaeology, Palynology, Pedoanthracology and linguistic research in order to address the question of the beginning of alpine farming in the Alps. With the discovery of remains from alpine dairy huts, archaeological studies show that there is definite proof of alpine farming beginning in the Bronze Age (2200–800 BC) in different parts of the Alps. Archaeological and palynological data as well as linguistic findings from many different studies and study areas arrived at the same conclusions. Palynological studies found indicators for high-altitude pasture use even earlier, beginning from 4500 BC. The exact type or intensity of this pasture use though remains unclear. In order to confirm these findings, more archaeological research of these areas would be necessary.

Land-use history and habitat age are considered highly important regarding the species composition and conservation value of, e.g., grasslands and forests. Ancient sites, i.e., sites with a continuous habitat history, are thought to be richer in rare, threatened, and more specialised species than recent sites. However, species composition is also affected by abiotic and ecological factors such as soil and climate parameters, and further analyses are needed to determine whether history or other factors are more important. In previous studies carried out at two study areas in the German Jurassic mountains, land-use history was identified as the main factor shaping vascular plant composition of ancient vs. recent grassland sites. Ancient was defined that there was a continuous grazing history since at least 200 years, while recent grasslands have developed from arable fields considerably after this time. Many species such as arable weeds and crop species, being residuals of the former land-use type, were found in the recent grasslands. In the present study, we assessed the terricolous cryptogam vegetation (bryophytes, lichens, macromycetes) of the same ancient and recent sites regarding species composition, occurrence of endangered species, and the applicability of the indicator species concept and Ellenberg indicator values. Although land-use history was the strongest explaining factor regarding vascular plant vegetation in previous studies, cryptogam vegetation did not reflect this signal. Species numbers and Ellenberg indicator values were quite similar in ancient and recent grasslands but we could, nevertheless, identify indicator species for both grassland types, with Rhytidium rugosum, Cladonia furcata ssp. subrangiformis and Hygrocybe persistens as strongest indicators of ancient grasslands, and Rhytidiadelphus squarrosus as strongest indicator of recent grasslands. When we compared the results for vascular plant and cryptogam vegetation we found that land-use history seems to have much less influence on the composition of the cryptogam vegetation in grasslands, given a certain time of recreation and succession. This is very likely due to an enhanced dispersability of cryptogam propagules. We found rare and endangered cryptogam species in ancient and recent sites; therefore, species maintenance measures can be worthwile in both grassland types.

Reassembling ecological communities and rebuilding habitats through active restoration treatments requires curating the selection of plant species to use in seeding and planting mixes. Ideally, these mixes should be assembled based on attributes that support ecosystem function and services, promote plant and animal species interactions and ecological networks in restoration while balancing project constraints. Despite these critical considerations, it is common for species mixes to be selected opportunistically. Reframing the selection of seed mixes for restoration around ecological objectives is essential for success but accessible methods and tools are needed to support this effort.We developed a framework to optimize species seed mixes based on prioritizing plant species attributes to best support different objectives for ecosystem functions, services, and trophic relationships such as pollination, seed dispersal, and herbivory. We compared results to approaches where plant species are selected to represent plant taxonomic richness, dominant species, and at random. We tested our framework for 176 plant species found in European alpine grasslands and identified 163 associated attributes affiliated to trophic relationships, ecosystem functions, and services.In all cases, trophic relationships, ecosystem functions, and services can be captured more efficiently through objective-based prioritization using the functional identity of plant species. Solutions (plant species lists) can be compared quantitatively, in terms of costs, species, or objectives. We confirm that a random draw of plant species from the regional plant species pool cannot be assumed to support other trophic groups and ecosystem functions and services.Synthesis and Applications. Our framework is presented as a proof of concept to help restoration practitioners better apply quantitative decision–support to plant species selection in order to meet ecological restoration outcomes. Our approach may be tailored to any restoration initiative and habitat where seeding or planting mixes will be applied in active treatments. As global priority and resources are increasingly placed into restoration, this approach could be advanced to help make efficient decisions for many stages of the restoration process.

The abandonment of historical land‐use forms within forests, such as grazing or coppicing, and atmospheric nitrogen deposition, has led to an increasing overgrowth of forest gaps and canopy closure in forest ecosystems of Central Europe. From 1945 to 2015, 81% of the forest gaps greater than 150 m2 within the study area transitioned into a closed forest.This study investigated how the overgrowth process affects flower supply, flower visitors, and reproduction of Campanula species. Six native Campanula species with different light requirements were used as phytometers.The forest gaps in the studied area are a feature of the historical European cultural landscape. We compared large gaps caused by human activities, small gaps caused by habitat conditions, and closed forests. In eight blocked replicates, each with the three habitat categories, we recorded the flower cover and number of indigenous flowering species in the immediate surroundings, and, of six Campanula species, flower visitors and seed production.Forest gaps and their size positively affected the number of flowering plant species in the surrounding area, the number of all flower visitor groups, and the number of seeds produced by all six Campanula species. Flower cover in the surrounding area was higher in large gaps, but there was no difference between small gaps and closed forests. Among flower visitors, small bees varied the most between the three habitat categories, and flies varied the least. The effect on the number of seeds produced was particularly strong for three light‐demanding Campanula species.The overgrowth of forest gaps negatively affected flower supply, flower‐visiting insects, and seed sets of six Campanula species. Forest gaps should be managed to maintain the reproduction of open forest plants and their pollinators., Nitrogen deposition and abandonment of extensive forms of land use lead to an overgrowth of forest gaps. This study compared the flower supply, flower visitors, and seed set of six Campanula species of large and small forest gaps and closed forest. The overgrowth of forest gaps negatively affected flower supply, flower‐visiting insects, and seed sets of six Campanula species. (The photograph in the center shows Hoplitis mitis visiting Campanula persicifolia and seeds of C. persicifolia.)

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. ispartof: NEW PHYTOLOGIST vol:229 issue:6 pages:3573-3586 ispartof: location:England status: published

Aims Persistence plays a key role in both plant population and community dynamics as it allows withstanding temporarily habitat conditions unsuitable for growth and reproduction. Plants can exploit three different strategies to persist in situ (building soil seed banks, increasing adult longevity and/or clonal reproduction), the dominance of which in a community might vary along environmental gradients. Yet, their relative role in plant persistence has never been investigated. Location The Bavarian Alps, Germany. Methods We collected data on seed soil persistence, adult longevity and clonality for 290 species occurring in 18 grasslands located along an elevational gradient of 1,000 metres and examined their contribution to persistence in a community. Linear models were used to estimate the relationship between elevation and these persistence strategies. Results We found that dominance of a certain persistence strategy varied depending on environmental variability. Specifically, persistence in lowlands was mainly achieved by persistent and dense soil seed banks along with extended clonal growth (larger spread distances and higher number of offspring). Contrastingly, the main persistence strategy in the alpine communities was increased adult longevity. Conclusions The changes in relative contribution of each strategy to community persistence along the elevational gradient are interrelated suggesting a trade-off among them. We conclude that this trade-off plays an important role in species co-existence and community assembly, and might be useful to understand vegetation dynamics under ongoing climate change and improve restoration efforts of upland ecosystems.