Relatively open vegetation landscapes promoted early Pleistocene hominin evolution

As cities grapple with climate risks and biodiversity decline, international policy frameworks increasingly position urban nature as a critical ally in building resilient and sustainable urban areas. The European Union's Biodiversity Strategy 2030, for example, calls on European cities to develop and implement ambitious 'Urban Nature Plans' (formerly 'Urban Greening Plans'). In this paper, we introduce Urban Nature Plans as a strategic planning tool and discuss potential pathways to their implementation, using Germany as an example. With more than 650 cities of diff erent sizes and with diff erent administrative capacities – many of which already have planning instruments sharing features with Urban Nature Plans – Germany provides a useful case to discuss the implications of introducing this new strategic approach. To contribute to the international discourse on Urban Nature Plans and wider discussions around integrated biodiversity and climate planning, we summarize findings from a German research and development project. The project included, among other aspects, a dialogue process with government organizations, professional associations, non-governmental environmental organizations, municipal representatives and other actors. During this process, it became evident that Urban Nature Plans should be understood as ambitious, integrated, and flexible strategies which are adaptable to local needs and capacities. A clear commitment from local politicians is seen to play a critical role in increasing the amount and quality of urban nature, promoting the participation of diff erent stakeholder groups and accelerating the implementation of related measures. While existing planning instruments such as open space plans and landscape plans can ful fil many of the Urban Nature Plan requirements, a new and targeted strategy could address gaps and shortcomings of existing approaches, such as a lack of integrated working methods, co-design approaches or monitoring mechanisms. However, the findings also highlight that cities have diff erent capacities for strategic planning and the implementation of measures, and that Urban Nature Plans need to correspond to these diff erences if they are to be more than a tool for larger cities and existing champions in sustainability across Europe.

This paper provides a comprehensive analysis of the 2025 Altai film The Wolves (Волки), directed by Mikhail Kulunakov, positioning it within the burgeoning phenomenon of Russian regional cinema. In recent years, cinematic production from Russia's constituent republics, such as Yakutia (Sakha) and Altai, has garnered international acclaim for its unique cultural and linguistic specificity, operating outside the mainstream Moscow-St. Petersburg film industry. Based on a true 19th-century story, The Wolves depicts a personal conflict originating from a traditional bride-kidnapping that escalates into a brutal, community-wide vendetta. Through a multi-faceted approach combining narrative, aesthetic, and comparative analysis, this paper examines how the film portrays the cyclical nature of violence and revenge within the cultural context of Altai. The study argues that The Wolves embodies key characteristics of Russian regional cinema, including a profound connection to ethnic roots and the mythical power of nature. A central focus is placed on the film's core paradox of “fake wolves versus real violence,” where humans mimic the sounds and behaviors of wolves to perpetrate a brutality that surpasses that of the actual animals. The Altai Taiga is analyzed not as a mere backdrop but as a “silent arbiter”—a living entity that impassively witnesses human devolution. The aesthetic analysis highlights Olga Kulunakova's cinematography, which utilizes majestic long shots, natural lighting, and a stark contrast between vast, open landscapes and claustrophobic interiors to visualize the film's thematic tensions. Furthermore, by comparing The Wolves with the works of acclaimed Yakutian director Dmitry Davydov, the paper identifies shared thematic concerns—such as violence in isolated communities—while also distinguishing its epic, historical-drama aesthetic from Davydov's documentary-style realism. In conclusion, this paper posits that The Wolves is a landmark film that not only artistically represents the unique cultural identity of Altai but also exemplifies the growing artistic and thematic diversity of the Russian regional cinema movement, offering a profound meditation on the universal themes of violence, honor, and the ambiguous boundary between humanity and wildness.

The Dutch Land Goat (Nederlandse Landgeit) is one of Europe's oldest goat breeds, originally developed as a resilient smallholder's livestock in the Netherlands. Today, it plays an increasingly important role in conservation grazing projects, particularly on small-scale and structurally complex sites where traditional grazing animals such as sheep are of limited use. The breed is characterized by high robustness, adaptability, and a distinct browsing behavior with a preference for shrubs and woody vegetation. This makes it especially effective in controlling invasive or competitive plant species such as blackberry (Rubus sect. Rubus), holly (Ilex), and late-flowering black cherry (Prunus serotina).By browsing and bark-stripping, Dutch Land Goats contribute not only to maintaining open cultural landscapes but also to a long-term reduction of undesirable vegetation. Their relatively low tendency to jump, protective long coat, and strong keeper orientation facilitate practical management. Nevertheless, small-scale projects and specific conservation goals require close monitoring of health and nutrition. Key aspects include adequate mineral supplementation (e. g., selenium), claw health on moist soils, and prudent parasite management to minimize adverse effects on dung fauna.Experiences at Tierpark Nordhorn demonstrate that male groups are particularly suitable for seasonal use, as they remain more manageable than females with offspring. Mobile handling and restraining systems adapted to the goats' physical abilities are essential, while regular interactions at feeding and watering points enhance trust and facilitate veterinary care.Dutch Land Goats thus represent a valuable component of conservation grazing systems. Their specialized browsing behavior enables effective management of shrub-encroached habitats on small areas. Moreover, as a rare heritage livestock breed, they provide an important contribution to the preservation of genetic diversity.

The distribution of adipose depots in different body parts affects pig production value and human health, governed by complex epigenomic mechanisms. Limited studies on pig adipose depots have hindered the genetic improvement of fat-related economic traits and their biomedical applications. To address this issue, we generated epigenomic maps for backfat, belly fat, groin fat, and intermuscular fat (IMF) in Meishan pigs, integrating ChIP-seq, ATAC-seq, RNA-seq, Hi–C, and public whole-genome sequencing data. Our results reveal that belly/backfat share similar chromatin states, while groin fat/IMF exhibit distinct H3K27ac modification, super-enhancer (SE) dynamics, and open chromatin landscapes compared to belly/backfat. The spatially specific expressions of adipogenic transcription factors (TFs), such as lipid synthesis-related TFs PPARA and SOX6, which are highly expressed in back/belly fat, and adipocyte differentiation TF KLF4 was driven by a groin fat specific SE, underlie these chromatin state disparities. These results also suggest enhanced lipid synthesis in belly/backfat and adipocyte differentiation in groin fat. Moreover, candidate functional variants identified in IMF-gained H3K27ac peaks are primarily associated with meat quality traits. Genes linked to pig backfat thickness may also serve as candidate genes for human obesity due to the conserved cis-regulatory elements and gene expression patterns between humans and pigs. Overall, our epigenomic landscape enhances understanding of adipose depot regulation in mammals, facilitating cross-species insights and precision breeding.Supplementary InformationThe online version contains supplementary material available at 10.1186/s13578-025-01513-8.

Vehicle-mounted cameras reveal negative impact of the Fukushima Daiichi nuclear power plant accident on large-bodied bird abundance via paddy field abandonment.

ABSTRACT Russia has been involved in Africa since Tsarist times, but its involvement has ebbed and flowed. Nonetheless, the idea of Africa as an open political landscape in which Russia could act as a great power has been relatively persistent, often in connection with the domestic resurfacing of civilisational discourses on Russian exceptionalism. Recently, there has been much concern in the West about the expansion of Russian projection in Africa, with some suggesting that Moscow is establishing a sphere of influence. Our analysis of Russia’s geopolitical imagination and its subsequent practices does not lead us to conclude that a deliberate attempt to establish a genuine sphere of influence is underway. Instead, we see an opportunistic endeavour to assert and shape a geographically continuous sphere of interest (Russian: sfera interesov), negotiating with local partners forms of collaboration that reflect and prompt change in the international system.

Zygotic genome activation is tightly associated with the modulation of chromatin accessibility via maternal transcription factors. Understanding how chromatin accessibility is established and identifying key maternal regulators are crucial to comprehending this process. Here, by developing CANTAC-seq, we generate a genome-wide map of accessible chromatin of early Xenopus tropicalis embryos and find that the open chromatin landscape is progressively established at cis-regulatory elements during zygotic genome activation. Based on the motif analysis and perturbation experiments, we demonstrate that E2f1 maintains a repressive chromatin environment and inhibits zygotic gene transcription before the mid-blastula transition. Moreover, we identify that Otx1, another maternal transcriptional activator, coordinates with E2f1 in regulating chromatin accessibility and zygotic genome activation. Together, E2f1 and Otx1 determine the timely expression of a subset of genes required for zygotic gene transcription and germ layer differentiation.

Cultivation by humans is the primary mode of introduction for naturalized plants and an important driver of naturalization, a critical step in the invasion process. Historical records of cultivated plants can represent introduced species pools and propagule pressure, allowing for tests of how species' traits and environmental context affect naturalization while accounting for human influence. Ruderal traits, which generally promote naturalization, may not be universally advantageous across closed versus open landscapes (forest versus grassland/shrubland) or different agricultural land use conversion types, though such context dependence has not yet been demonstrated at a broad scale. We analyzed the naturalization of 3949 cultivated ornamental non‐native plant taxa that were for sale in nursery and seed catalogs in the conterminous United States during a period over 200 years to test for context dependence between traits associated with ruderality (short lifespan, shade intolerance, and self‐compatibility) and estimates of historical forest/grassland cover and agricultural land use change. We found that present‐day naturalization was closely tied to longer cultivation duration and greater cultivation extent. While ruderal traits tended to promote naturalization, perennial lifespan and shade tolerance favored naturalization in US states with higher forest cover, which is consistent with an alternative invasion strategy in closed‐canopy systems. Land use conversion to pasture and succession of abandoned agricultural land promoted naturalization of disturbance‐adapted plants in both forest and grassland landscapes. Our results emphasize the central role of cultivation in plant invasion and provide spatially and temporally extensive evidence that, while ruderal traits are important predictors of naturalization, they are dependent on the landscape context into which plants are introduced. Our work demonstrates the importance of integrating historical cultivation and land use/cover data for a nuanced understanding of the ecological factors that drive plant naturalization.

The Green Belt policy has historically played a crucial role in containing urban sprawl and preserving open landscapes in the UK. However, it has come under increasing pressure from rising housing demands and the push for economic growth in recent decades. This study uses the fictional High Vale Film Studio Project as a case study to explore how urban expansion can be reconciled with environmental conservation and social welfare. Based on a literature review and multi-stakeholder analysis, this paper shows that sustainable outcomes are dependent on conditional and adaptive strategies, such as robust ecological safeguards, meaningful public involvement, and policy innovation. These will allow for careful land-use adjustments. The findings provide a practical framework for reforming Green Belt policy which can balance development needs with long-term sustainability. It also provides insights applicable to similar peri-urban contexts facing growth pressures.

The strength of fragile grounds lies in their unfinished features, their ‘ruined’ patina and thus in their being weak, porous and in continuous deformation; but they are also open and complex landscapes, available to change repeatedly. The physical and cultural vulnerability of fragile grounds makes it difficult for society to imagine a coexistence with it. However, the accidental component, meant as an occasional but at the same time creative relationship with the landscape, is both the intrinsic dynamism of fragile grounds and the different possibilities of using the ordinary space of everyday life. This contribution investigates the issue from a qualitative and no longer just quantitative point of view, through the discipline of architecture. Referring to the methodological aspect, new variables are identified, as well as some interpretative and operative devices, which flow into the project in specific actions for the valorization and enhancement of fragile grounds in stratified contexts of historical-archaeological and landscape relevance. This approach makes it possible to deepen the concept of inter-spaces, which, starting from an intrinsic condition of vulnerability of a ‘middle ground’, creates the conditions for a project of interrelation and contributes to the development of a significant role of architecture in contexts exposed to risks.

Roads are among the most widespread anthropogenic structures, and their presence can impact biodiversity in surrounding landscapes through disturbance and collision risk, particularly when traffic volumes are high. However, the impact of roads with low traffic volumes in open landscapes is much less clear. In the open landscapes of lowland Iceland, road traffic is still relatively low but increasing, and the surrounding landscapes support internationally important populations of several breeding wader species. Here, we used transect counts perpendicular to low‐traffic (≤ 15 000 vehicles day −1 ) roads across the lowlands of southern Iceland to quantify variation in the densities of ground‐nesting birds with distance from roads, and to assess how far from the roads any such effects extended. The total abundance of birds increased significantly by 6% per 50 m interval from roads, and densities within 200 m of roads were ~ 20% lower than densities between 200 and 400 m from roads. Four species – whimbrel Numenius phaeopus , golden plover Pluvialis apricaria , dunlin Calidris alpina and meadow pipit Anthus pratensis – were found in significantly lower densities closer to roads, while four – black‐tailed godwit Limosa limosa , redshank Tringa totanus , snipe Gallinago gallinago and redwing Turdus iliacus – showed no change with distance from roads. Redwing was found in higher densities, and dunlin in lower densities, surrounding roads with higher traffic volumes. As approximately 20% of lowland Iceland is within 200 m of roads, the impact of roads on the overall abundance of ground‐nesting birds could be substantial. The results show that even relatively low‐traffic roads can have a significant impact on adjacent wildlife populations. Road construction, along with other anthropogenic structures, has been shown to have negative effects on bird abundance, and identifying areas for protection from such developments may be the most effective approach to reducing human impacts on the internationally important wildlife of lowland Iceland.

Abstract Declining forest biodiversity has increased focus on forest conservation and restoration. Many efforts to conserve and restore tree cover focus on the local scale, but their outcomes are frequently modulated by landscape context. While the diversity and composition of communities is strongly driven by local‐scale canopy cover, landscape‐scale habitat characteristics affect dispersal pathways and determine the species pool available for colonization of local patches. Moreover, local and landscape‐scale habitat attributes vary with elevation, but how their effects on biodiversity change with elevation remains poorly understood. We examined how local canopy cover affects forest biodiversity, and how these effects are modulated by the amount of total and disturbed forest available in the surrounding landscape along an elevational gradient. We used remote sensing and multi‐taxa biodiversity data covering plants, aculeata, moths, beetles and birds (a total of 2319 species) across 150 plots in naturally developing forests in a forest‐rich region in the northern European Alps. We calculated multidiversity across all species and for three habitat affinity guilds (forest, mixed, open‐habitat) to test for differences based on varying habitat associations. Local canopy cover negatively affected multidiversity, with the weakest effect observed for forest species. An increasing amount of forest in the surrounding landscape amplified this negative effect, while an increasing amount of disturbed forest reduced it. The negative effect of local canopy cover on multidiversity weakened with elevation and became neutral across all guilds close to the tree line. Our findings highlight that disturbances promote forest biodiversity via two fundamental pathways: reducing local canopy cover and creating a more open and diverse landscape context. Moreover, the effects of canopy cover on forest biodiversity are modulated by environmental conditions that change with elevation. Synthesis and applications . Conservation and restoration efforts should consider landscape context more explicitly when planning specific management measures. Our results suggest that canopy openings benefit biodiversity especially in landscapes with high forest cover and in low elevation areas, while conserving and re‐establishing tree cover is important in landscapes with low forest cover and close to the upper tree line.

This article presents an open schooling landscape education paradigm implemented in secondary education in Greece. Although Geography is the core subject addressing Environment and Sustainability Education (ESE) in the national curriculum, it remains significantly marginalized within the educational system. The article identifies and discusses the key dimensions of this systemic marginalization in order to inform the international academic community. Likewise, ESE as a whole is undervalued and insufficiently integrated into educational practice, a condition that is largely attributable to the diminished status of Geography itself. The paper argues that Geography matters and that it can be revitalized through landscape-based open schooling. Within this framework, landscape is introduced as a central geographical concept that enables a holistic, experiential, and applied understanding of space and environment. Twenty-eight students participated in this participatory qualitative research, employing methods including brainstorming, flipped classroom, collaborative group work, and project-based learning. Students were actively engaged both in classroom dialogue and as “young researchers” in the field, generating proposals for sustainable rural landscape management. Building on this framework, the study explores how open schooling landscape education, can cultivate scientific literacy, enhance environmental awareness, and strenthen student’s engagement with sustainability through experiential, place-based, and problem-oriented learning. The landscape concept served as a multidisciplinary entry point, bridging geography, science, and education. Findings show that the approach promotes both theoretical and practical learning, nurtures key skills and attitudes, strengthens collaboration among schools and communities, and supports the science capital and spatial awareness. It further lays the groundwork for active citizenship and more responsible human–landscape relations. Integrating open schooling landscape education into the formal curriculum is strongly advocated.

Abstract. Mobile multicam photogrammetry systems offer a number of advantages over other 3d imaging modalities, including rapid acquisition speed, low cost sensors, and the availability of mature open-source software systems for photogrammetric alignment and reconstruction.. Through a number of case studies we compare the Looq, a home-grown GPS enabled multi-cam mobile mapping system, to colocated models from terrestrial photogrammetry, aerial photogrammetry, two terrestrial LiDAR systems, and two SLAM LiDAR systems. We evaluate these systems across varied urban and natural environments, in GPS enabled and denied environments, tight interior spaces, and open landscapes in Europe and North America. The comparative performance of these systems is described qualitatively and quantitatively.

The Late Pleistocene was a time of global megafaunal extinctions that were particularly severe in North America. The continent lost many mammal taxa, but the validity of several remains ambiguous, including a high proportion of Cervidae taxa. Torontoceros hypogaeus is represented by a single specimen (ROMM75974) discovered in 1976 during excavation work for the Toronto subway in Canada. The species was described based on its unique antler morphology, but the variable nature of that trait and the species near absence in the fossil record leads to uncertainty concerning its systematic relationships. We used ancient DNA to clarify the taxonomic relationship and evolutionary history of T. hypogaeus. We performed mitochondrial and whole genome analyses with related cervids and showed that ROMM75974 has a close affinity, but relatively high divergence from Odocoileus sister species. While some ambiguity remains, ROMM75974 could represent a distinct Odocoileus species to be included in the list of extinct North American taxa. This unique population was likely adapted to open landscape, which was rapidly replaced with dense woodland in this region at the end of the Pleistocene, highlighting the role of climate change in the extinction of megafauna biodiversity at the end of the ice age.

ABSTRACTThe Brazilian Cerrado is a continental‐wide biodiversity hotspot and the most species‐rich savanna ecosystem in the world. The main aspect characterising this biodiversity is that the landscape is arranged as an intricate mosaic of different plant formations, including grasslands, savannas, and forests, each harbouring distinct but interconnected communities. Seasonality and natural fires are key and ancient natural factors in the biome, with organisms showing many adaptations. The Cerrado is also home to millions of people, and the essential ecosystem services provided for agricultural production make it one of the world's major crop regions. However, it has undergone intense destruction in the last decades, with conservation concerns historically overshadowed by the neighbouring Amazonia and Atlantic Forest biomes. Considering the importance of pollination and plant reproduction for maintaining terrestrial ecosystems, we synthesise the known information for the Cerrado as an illustrative example that could be applied to other megadiverse ecosystems worldwide. Although apomixis (asexual seed formation) and self‐pollination mechanisms occur to a lesser extent, most plants in the Cerrado require biotic pollination. For instance, this is the case for some dioecious and monoecious species. However, the majority of plants have bisexual flowers, with the frequency of self‐incompatibility increasing towards denser plant formations such as forests, illustrating differences in dependency on pollination across habitats. Many Cerrado plants adopt strategies favouring outcrossing, including distyly, enantiostyly, heteranthery, and dichogamy. Although plant–pollinator interaction networks are mostly generalised, the pollinators are organised into guilds, with bees pollinating most plants and using several resources. Other common guilds include beetles, moths, hummingbirds, and bats. Importantly, flowering phenology peaks across plant formations at different times of the year, creating habitat complementarity across the vegetation mosaic that continuously sustains transiting pollinators. Thus, the interaction between plants and pollinators connects and is sustained by landscape complexity, which should be regarded as essential for ecosystem conservation. In this context, periodic fires that trigger massive flowering and promote biomass reduction are an essential natural disturbance that maintains the diversity of open landscapes. The interdependence of plants and pollinators in the face of the ongoing destruction of the Cerrado adds another challenge for its conservation, and highlights the necessity for conserving complementary habitats at the landscape level. While forest formations are granted protection by law, these alone are insufficient to maintain high pollinator diversity, with potential cascading effects on the ecosystem services they provide and requiring the maintenance of the neglected grasslands and savannas. Thus, the simultaneous conservation and restoration of the mosaic plant formations across the landscape will be crucial for the future of the Cerrado.

Abstract This study evaluates the potential of horses in Mediterranean forest management through their feeding behaviours. While traditionally overshadowed by livestock like sheep and goats, horses demonstrate significant adaptability, making them possible effective agents for reducing fuel loads and maintaining open landscapes. The findings highlight distinct feeding strategies among horse types. Feral Przewalski’s horses, in a mosaic landscape, with patches of forest, scrub and open grassland mostly consume graminoids throughout the year, maintaining open grasslands with minimal impact on woody vegetation. Rustic breeds, such as the Pottoka, adapt over extended grazing periods by initially consuming highly flammable graminoids and transitioning to woody plants as grasses deplete, effectively managing both fine and coarse fuels. Crossbred horses, under short-term, high-intensity grazing with supplemental feeding, rapidly shift from fine fuels to woody plants. Diet was quantified from 50 fresh faecal samples using micro-histological analysis and compositional Dirichlet regression. Although the study does not directly quantify vegetation changes or reductions in flammable biomass, the observed dietary shifts—particularly the increased consumption of woody species—highlight horses' potential role in understory management. These results suggest that different horse types respond flexibly to forage availability and may be useful in integrated strategies for reducing fire risk, especially when combined with other herbivores or active management practices. Overall, horses present a flexible and natural tool for forest management, with preliminary and exploratory evidence suggesting that they may contribute to fuel reduction through adaptive foraging behaviour, although further studies are needed to confirm vegetation impacts. Their use complements traditional fire prevention techniques and promotes the semi-freedom of horses, supporting their inclusion in sustainable landscape management.

Sexual conflict over mating has been documented in many species, both in the field and in experimental studies. In pond damselflies (family Coenagrionidae), sexual conflict maintains female-limited color polymorphisms, with one female morph typically being a male mimic. However, it is not known whether sexual conflict can also explain the evolutionary origin of novel female morphs, and if so, what ecological factors play a role in this macroevolutionary transition, by modulating the strength of the conflict. Here, we use phylogenetic comparative methods to show that female color polymorphisms are more likely to evolve in temperate regions and open landscapes, whereas region and habitat shifts are independent of female-color states. We also show that these macroecological patterns are mediated by population densities at breeding sites. Temperate and open habitats are associated with female-polymorphic lineages because they harbor higher densities of adults, promoting more frequent encounters between females and males. Finally, we found that female-limited polymorphisms typically evolve from sexually dimorphic ancestors through the addition of a male-like female morph, consistent with the hypothesis of selection for male mimicry. We conclude that female color polymorphisms evolve in a predictable fashion and are likely driven by ecological conditions that increase the rate of premating interactions and thus the intensity of sexual conflict.

Scarturus williamsi (Williams' jerboa) is a medium-sized, semi-fossorial rodent endemic to steppe ecosystems across Anatolia, Iran, and Azerbaijan, with specialized habitat requirements in semi-arid continental environments. This study integrates a mitochondrial DNA analysis with species distribution modeling to assess the species' evolutionary structure and vulnerability to future climate change. The phylogeographic analysis and species distribution modeling reveal the evolutionary history and climate vulnerability of Scarturus williamsi across Anatolia and adjacent regions. The mitochondrial DNA analysis of 98 individuals demonstrates exceptional haplotype diversity (Hd = 0.9896), with 90 unique haplotypes and complete regional isolation, indicating pronounced population structuring across five evolutionary lineages: Central Anatolia, Eastern Anatolia, Aegean, Black Sea, and Azerbaijan-Iran. The Iran-Azerbaijan lineage exhibits the deepest evolutionary divergence, while Eastern Anatolia functions as the primary Anatolian refugium and Central Anatolia as the secondary refugial center. The strong isolation by distance (r = 0.735, p < 0.001) across ~2500 km explains 54.0% of the genetic variation, with the hierarchical structure reflecting greater Iran-Turkey isolation than intra-Turkish differentiation. The species distribution modeling identifies the Mean Temperature of Driest Quarter (bio9) and the Mean Diurnal Range (bio2) as primary habitat determinants, with bimodal preferences reflecting highland versus steppe adaptations. Climate projections reveal severe vulnerability with habitat losses of 63.69-98.41% by 2081-2100 across emission scenarios. SSP3-7.0 represents the most catastrophic scenario, with a severe habitat reduction (98.41% loss), while even optimistic scenarios (SSP1-2.6) project a 60-70% habitat loss. All scenarios show accelerating degradation through mid-century, with the steepest losses occurring between 2041 and 2080. Projected eastward shifts face constraints from the Anatolian Diagonal, limiting the climate tracking capacity. Despite occupying open landscapes, S. williamsi exhibits exceptional sensitivity to climate change, with Anatolian refugial areas representing critical diversity centers facing substantial degradation. Results provide baseline genetic structure and climate vulnerability information for understanding climate impacts on S. williamsi and Irano-Anatolian steppe fauna.