Unforeseen plant phenotypic diversity in a dry and grazed world.

Earth harbours an extraordinary plant phenotypic diversity1 that is at risk from ongoing global changes2,3. However, it remains unknown how increasing aridity and livestock grazing pressure-two major drivers of global change4-6-shape the trait covariation that underlies plant phenotypic diversity1,7. Here we assessed how covariation among 20 chemical and morphological traits responds to aridity and grazing pressure within global drylands. Our analysis involved 133,769 trait measurements spanning 1,347 observations of 301 perennial plant species surveyed across 326 plots from 6 continents. Crossing an aridity threshold of approximately 0.7 (close to the transition between semi-arid and arid zones) led to an unexpected 88% increase in trait diversity. This threshold appeared in the presence of grazers, and moved toward lower aridity levels with increasing grazing pressure. Moreover, 57% of observed trait diversity occurred only in the most arid and grazed drylands, highlighting the phenotypic uniqueness of these extreme environments. Our work indicates that drylands act as a global reservoir of plant phenotypic diversity and challenge the pervasive view that harsh environmental conditions reduce plant trait diversity8-10. They also highlight that many alternative strategies may enable plants to cope with increases in environmental stress induced by climate change and land-use intensification.

Ecological restoration and biodiversity-friendly management of urban grasslands - A global review on the current state of knowledge.

In the face of the global biodiversity decline, ecological restoration measures to actively enhance urban biodiversity and options for biodiversity-friendly greenspace management are high on the agenda of many governments and city administrations. This review aims to summarize and advance the current knowledge on urban grassland restoration by synthesizing research findings on restoration approaches and biodiversity-friendly management measures globally. Indeed, we found restoration approaches to be generally effective in increasing biodiversity; yet, there were variations in the outcomes due to the difference in soil disturbance methods, management regimes, the set of species introduced to a site, and the specific local setting. Based on the reviewed studies, we formulated recommendations for maximizing restoration success of urban grasslands through: i) creating a network of heterogeneous urban greenspaces and enhancing connectivity between them; ii) maintaining the spontaneous vegetation in vacant lots and wasteland sites that can provide habitats for various invertebrate species; iii) evaluating actual soil conditions, soil seed bank, and seed rain before restoration efforts take place since these seed sources could considerably affect the restoration outcomes, iv) preserving nutrient-poor conditions in urban greenspaces instead of introducing nutrient-rich topsoil; v) shifting to less intensive, biodiversity-friendly management in urban greenspaces by reducing mowing frequency and avoiding the use of chemicals; and vi) utilizing native dry grassland species for climate adaptation without irrigation. We further identified knowledge gaps regarding i) city-scale and regional-scale effects of restoration, ii) effects of interventions on multiple taxa and multiple ecosystem services, iii) restoration in small versus mega-cities, and iv) in the global south. These gaps should be addressed in future studies for making general guidelines for urban grassland restoration broadly applicable.

Contribution of cultural heritage values to steppe conservation on ancient burial mounds of Eurasia.

Civilizations, including ancient ones, have shaped global ecosystems in many ways through coevolution of landscapes and humans. However, the cultural legacies of ancient and lost civilizations are rarely considered in the conservation of the Eurasian steppe biome. We used a data set containing more than 1000 records on localities, land cover, protection status, and cultural values related to ancient steppic burial mounds (kurgans); we evaluated how these iconic and widespread landmarks can contribute to grassland conservation in the Eurasian steppes, which is one of the most endangered biomes on Earth. Using Bayesian logistic generalized regressions and proportional odds logistic regressions, we examined the potential of mounds to preserve grasslands in landscapes with different levels of land-use transformation. We also compared the conservation potential of mounds inside and outside protected areas and assessed whether local cultural values support the maintenance of grasslands on them. Kurgans were of great importance in preserving grasslands in transformed landscapes outside protected areas, where they sometimes acted as habitat islands that contributed to habitat conservation and improved habitat connectivity. In addition to steep slopes hindering ploughing, when mounds had cultural value for local communities, the probability of grassland occurrence on kurgans almost doubled. Because the estimated number of steppic mounds is about 600,000 and similar historical features exist on all continents, our results may be applicable at a global level. Our results also suggested that an integrative socioecological approach in conservation might support the positive synergistic effects of conservation, landscape, and cultural values.

Contribución de los valores culturales para la conservación esteparia en los antiguos montículos funerarios de Eurasia Resumen Las civilizaciones modernas y antiguas han moldeado de muchas maneras los ecosistemas globales mediante la coevolución del paisaje y la humanidad. Sin embargo, pocas veces se considera el legado cultural de las civilizaciones perdidas o antiguas para la conservación del bioma de la estepa euroasiática. Usamos un conjunto de datos que contiene más de 1,000 registros de las localidades, cobertura del suelo, estado de protección y valores culturales relacionados con los antiguos montículos funerarios de esta estepa (kurgans). Después analizamos cómo estos símbolos icónicos y distribuidos extensamente pueden contribuir a la conservación de los pastizales en la estepa euroasiática, uno de los biomas en mayor peligro de extinción. Analizamos el potencial de conservación de los montículos en paisajes con diferentes niveles de transformación en el uso de suelo mediante regresiones logísticas generalizadas bayesianas y regresiones logísticas de probabilidades proporcionales. También comparamos el potencial de conservación de los montículos dentro y fuera de las áreas protegidas y evaluamos si los valores culturales locales conservan los pastizales dentro de estas mismas áreas. Los kurgans fueron de gran importancia para la conservación de los pastizales en los paisajes transformados ubicados fuera de las áreas protegidas, en donde llegaron a fungir como hábitats aislados que contribuyeron a la conservación y conectividad del hábitat. Además de que las pendientes pronunciadas impiden el arado, cuando los montículos contaban con valor cultural para las comunidades locales, la probabilidad de que el pastizal se ubicara sobre un kurgan casi se duplicó. Ya que se estima que el número de montículos esteparios ronda los 6,000 y que rasgos históricos similares existen en todos los continentes, nuestros resultados pueden aplicarse a nivel global. Nuestros resultados también sugieren que una estrategia socio-ecológica integradora para la conservación podría respaldar los efectos sinérgicos positivos de la conservación, el paisaje y los valores culturales.

Better safe than sorry - Understanding the attitude and habits of drivers can help mitigating animal-vehicle collisions.

The rapidly growing global road networks put serious pressures on terrestrial ecosystems and increase the number and severity of human-wildlife conflicts, which in most cases manifest in animal-vehicle collisions (AVCs). AVCs pose serious problems both for biodiversity conservation and traffic safety: each year, millions of vertebrates are roadkilled globally and the related economic damage is also substantial. For a comprehensive understanding of factors influencing AVC it is essential to explore the human factor, that is, the habits and attitude of drivers; however, to date, comprehensive surveys are lacking on this topic. Here we addressed this knowledge gap and surveyed the habits of drivers and their experience and attitude towards AVCs by a comprehensive questionnaire covering a large geographical area and involving a large number of respondents (1942 completed questionnaires). We aimed to reveal how driving habits affect the chance of AVC, and explored the attitude of the drivers regarding AVC. We found that the number of lifetime AVC cases was higher for male drivers, for those who drove longer distances per year, had more driven years, used country roads or drove large vehicles. Our results showed that almost half of the drivers surveyed had experienced at least one AVC in their lifetime. Drivers' attitudes towards the importance of nature conservation or traffic safety in the aspect of AVC, and fear of collision showed a significant correlation with experienced AVC cases. Drivers' opinions indicated that the most trusted and desired AVC prevention measures were physical objects such as fences and wildlife crossings. Our research provides guidelines for developing targeted initiatives in the future to increase awareness about the significance of AVC and target those drivers who are most vulnerable to AVC.

Hay transfer is a nature-based and sustainable solution for restoring grassland biodiversity.

As we enter the United Nations Decade on Ecosystem Restoration, there is an increasing need for finding nature-based solutions for the restoration of grasslands across the globe. Besides seed sowing, alternative restoration methods that build on locally available propagule sources, such as hay transfer, should also be considered and given high priority. The transfer of hay from a donor site to the restoration site serves a double aim by introducing target species and suppressing weeds. We tested the applicability of hay transfer in restoring species-rich grassland in a former cropland in Hungary, over six years of post-restoration monitoring from 2015 to 2020. We sampled the plant species composition of the three donor sites (target state of restoration) and the area surrounding the restoration site. We found that six year after restoration, a species-rich grassland developed with 42 successfully established specialist species that had a cover of 45%. Most specialist species became established from the hay from the second to the sixth year, but the surrounding areas also provided additional propagule sources. Among the eight analysed functional traits, seed mass of the transferred species was the best predictor of the establishment of specialists, and we found that small-seeded species had an establishment advantage in the first years after restoration. Our results suggest that hay transfer can be a suitable nature-based solution for local grassland restoration projects and its potential should be utilized especially in regions where suitable donor sites are present in sufficient quantity and quality.

Grassland restoration on linear landscape elements - comparing the effects of topsoil removal and topsoil transfer.

Artificial linear landscape elements, including roads, pipelines, and drainage channels, are main sources of global habitat fragmentation. Restoration of natural habitats on unused linear landscape elements can increase habitat quality and connectivity without interfering with agricultural or industrial development. Despite that topsoil removal and transfer are widely applied methods in restoration projects, up to our knowledge these were previously not compared in the same study system. To address this knowledge gap, we compared spontaneous vegetation recovery after the elimination of positive (embankments) and negative landscape scars (drainage channels) in lowland alkaline landscapes in South Hungary. The novelty of our study is that we compared the fine-scale and landscape-scale results of both methods. At the fine scale, we monitored the spontaneous vegetation development on the created open surfaces in the first, second and fourth year after restoration in 160 permanent plots per year. For characterizing the habitat changes on the landscape scale, we prepared habitat maps and assigned naturalness scores to each patch before and after the restoration activities. Both restoration methods resulted in a rapid vegetation recovery at the fine scale, progressing toward the reference state. In the topsoil removal treatment, a large part of the soil seed bank was removed; therefore, the colonization of the bare surface was a slower process. Seeds of halophytes, including the endemic and protected Suaeda pannonica, were probably present in the deeper soil layers, and these species became established in the restored surfaces, despite being absent in the surrounding vegetation. For restoring vegetation cover, topsoil transfer was a more rapid option; however, vegetation closure and competition by generalist species and weeds hampered the establishment of target species. The removal of the landscape scars by both methods made the sites accessible for grazing. At the landscape scale, the two methods had different effects: there was a slight increase in the habitat naturalness in the topsoil removal site, and a slight decrease in the topsoil transfer site because of weed encroachment. Spreading an upper layer of nutrient-poor soil with low amounts of weed seeds, direct propagule transfer, and targeted grazing regimes could enhance restoration success.

Meso-scale environmental heterogeneity drives plant trait distributions in fragmented dry grasslands.

Environmental heterogeneity shapes the patterns of resources and limiting factors and therefore can be an important driver of plant community composition through the selection of the most adaptive functional traits. In this study, we explored plant trait-environment relationships in environmentally heterogeneous microsite complexes at the meso-scale (few meters), and used ancient Bulgarian and Hungarian burial mounds covered by dry grasslands as a model habitat. We assessed within-site trait variability typical of certain microsites with different combinations of environmental parameters (mound slopes with different aspects, mound tops, and surrounding plain grasslands) using a dataset of 480 vegetation plots. For this we calculated community-weighted means (CWMs) and abundance models. We found that despite their small size, the vegetation on mounds was characterized by different sets of functional traits (higher canopy, higher level of clonality, and heavier seeds) compared to the plain grasslands. North-facing slopes with mild environmental conditions were characterized by perennial species with light seeds, short flowering period, and a high proportion of dwarf shrubs sharply contrasted from the plain grasslands and from the south-facing slopes and mound tops with harsh environmental conditions. Patterns predicted by CWMs and abundance models differed in the case of certain traits (perenniality, canopy height, and leaf dry matter content), suggesting that environmental factors do not necessarily affect trait optima directly, but influence them indirectly through correlating traits. Due to the large relative differences in environmental parameters, contrasts in trait composition among microsites were mostly consistent and independent from the macroclimate. Mounds with high environmental heterogeneity can considerably increase variability in plant functional traits and ecological strategies at the site and landscape levels. The large trait variation on topographically heterogeneous landscape features can increase community resilience against climate change or stochastic disturbances, which underlines their conservation importance.

Hotspots of biogeochemical activity linked to aridity and plant traits across global drylands.

Perennial plants create productive and biodiverse hotspots, known as fertile islands, beneath their canopies. These hotspots largely determine the structure and functioning of drylands worldwide. Despite their ubiquity, the factors controlling fertile islands under conditions of contrasting grazing by livestock, the most prevalent land use in drylands, remain virtually unknown. Here we evaluated the relative importance of grazing pressure and herbivore type, climate and plant functional traits on 24 soil physical and chemical attributes that represent proxies of key ecosystem services related to decomposition, soil fertility, and soil and water conservation. To do this, we conducted a standardized global survey of 288 plots at 88 sites in 25 countries worldwide. We show that aridity and plant traits are the major factors associated with the magnitude of plant effects on fertile islands in grazed drylands worldwide. Grazing pressure had little influence on the capacity of plants to support fertile islands. Taller and wider shrubs and grasses supported stronger island effects. Stable and functional soils tended to be linked to species-rich sites with taller plants. Together, our findings dispel the notion that grazing pressure or herbivore type are linked to the formation or intensification of fertile islands in drylands. Rather, our study suggests that changes in aridity, and processes that alter island identity and therefore plant traits, will have marked effects on how perennial plants support and maintain the functioning of drylands in a more arid and grazed world.

How to not trade water for carbon with tree planting in water-limited temperate biomes?

The most widespread nature-based solution for mitigating climate change is tree planting. When realized as forest restoration in historically forested biomes, it can efficiently contribute to the sequestration of atmospheric carbon and can also entail significant biodiversity and ecosystem service benefits. Conversely, tree planting in naturally open biomes can have adverse effects, of which water shortage due to increased evapotranspiration is among the most alarming ones. Here we assessed how soil texture affects the strength of the trade-off between tree cover and water balance in the forest-steppe biome, where the global pressure for afforestation is threatening with increasing tree cover above historical levels. Here we monitored vertical soil moisture dynamics in four stands in each of the most common forest types of lowland Hungary on well-drained, sandy (natural poplar groves, and Robinia and pine plantations) and on poorly drained, silty-clayey soils (natural oak stands and Robinia plantations), and neighboring grasslands. We found that forests on sand retain moisture in the topsoil (approx. 20 cm) throughout the year, but a thick dry layer develops below that during the vegetation period, significantly impeding groundwater recharge. Neighboring sandy grasslands showed an opposite pattern, with often dry topsoil but intact moisture reserves below, allowing deep percolation. In contrast, forests on silty-clayey soils did not desiccate lower soil layers compared neighboring grasslands, which in turn showed moisture patterns similar to sandy grasslands. We conclude that, in water-limited temperate biomes where landscape-wide water regime depends on deep percolation, soil texture should drive the spatial allocation of tree-based climate mitigation efforts. On sand, the establishment of new forests should be kept to a minimum and grassland restoration should be preferred. The trade-off between water and carbon is less pronounced on silty-clayey soils, making forest patches and wooded rangelands viable targets for both climate mitigation and ecosystem restoration.

Environmental heterogeneity increases the conservation value of small natural features in karst landscapes.

Local biodiversity hotspots are often located within regions where extreme and variable environmental - e.g., climatic and soil - conditions occur. These areas are conservation priorities. Although environmental heterogeneity is recognised as an important determinant of biodiversity, studies focusing on the effects of multiple environmental heterogeneity components in the same ecosystem are scarce. Here we investigate how topography and related microclimatic variables and soil properties may influence the biodiversity and conservation value of karst landscapes. Karst landscapes of the world contain millions of dolines (i.e. bowl- or funnel-shaped depressions) that may function as 'small natural features' with a disproportionately large role in maintaining biodiversity relative to their size. We assessed the diversity of microclimates, soils and vegetation and their relationships in six microhabitats (south-facing slopes, east-facing slopes, west-facing slopes, north-facing slopes and bottoms of dolines, and the adjacent plateau) for nine large dolines in a grassland ecosystem. Although there were remarkable differences among the conservation value of these microhabitats (e.g., representation of different species groups, presence of 'climate relicts'), each microhabitat had an important role in maintaining species that are rare or absent in other microhabitats in the landscape. We found that the studied dolines exhibited highly variable environmental conditions and promoted a high diversity of vegetation types with unique species composition, contributing to the topographic, climatic, soil, vegetation and land cover heterogeneity of karst landscapes. Therefore, our findings highlight that dolines may function as local biodiversity hotspots and have a crucial conservation importance. As dolines are widespread topographic features in many karst landscapes throughout the world, our results could be directly applied to other regions as well. An integrated approach is urgently needed to provide guidelines for landscape management, promoting the retention of the microhabitat diversity of small natural features for species vulnerable to climate change and/or various disturbances.