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Long-distance terrestrial migrations are imperiled globally. We determined both round-trip migration distances (straight-line measurements between migratory end points) and total annual movement (sum of the distances between successive relocations over a year) for a suite of large mammals that had potential for long-distance movements to test which species displayed the longest of both. We found that caribou likely do exhibit the longest terrestrial migrations on the planet, but, over the course of a year, gray wolves move the most. Our results were consistent with the trophic-level based hypothesis that predators would move more than their prey. Herbivores in low productivity environments moved more than herbivores in more productive habitats. We also found that larger members of the same guild moved less than smaller members, supporting the ‘gastro-centric’ hypothesis. A better understanding of migration and movements of large mammals should aid in their conservation by helping delineate conservation area boundaries and determine priority corridors for protection to preserve connectivity. The magnitude of the migrations and movements we documented should also provide guidance on the scale of conservation efforts required and assist conservation planning across agency and even national boundaries.

The non-equilibrium concept of rangeland dynamics predicts that the potential for grazing-induced degradation is low in rangelands with relatively variable precipitation. To date, evidence in support of the non-equilibrium concept has been inconsistent. Using a standardized protocol, including a newly developed global map of rainfall variability, we reviewed the incidence of degradation in relation to rainfall variability across 58 published studies. We distinguished between (1) zonal degradation (i.e., degradation independent of water and key resources), (2) degradation in the presence of key resources, and (3) degradation in the presence of water. For studies not affected by proximity to permanent water or key resources, we found strong support for the non-equilibrium concept for rangelands. Zonal degradation was absent at CV (coefficient of variation) values above 33%, which has been proposed as a critical threshold. Grazing degradation was almost entirely restricted to areas with relatively stable annual precipitation as expressed by a low CV, or to rangelands with key resources or water points nearby. To better understand rangeland dynamics, we recommend that future studies use globally comparable measures of degradation and rainfall variability. Our work underlines that rangelands with relatively stable rainfall patterns, and those with access to water or key resources, are potentially vulnerable to degradation. Grazing management in such areas should incorporate strategic rest periods. Such rest periods effectively mimic natural fluctuations in herbivore populations, which are a defining characteristic of non-degraded rangelands occurring under highly variable precipitation regimes.

While many animal species exhibit strong conspecific interactions, movement analyses of wildlife tracking datasets still largely focus on single individuals. Multi-individual wildlife tracking studies provide new opportunities to explore how individuals move relative to one another, but such datasets are frequently too sparse for the detailed, acceleration-based analytical methods typically employed in collective motion studies. Here, we address the methodological gap between wildlife tracking data and collective motion by developing a general method for quantifying movement correlation from sparsely sampled data. Unlike most existing techniques for studying the non-independence of individual movements with wildlife tracking data, our approach is derived from an analytically tractable stochastic model of correlated movement. Our approach partitions correlation into a deterministic tendency to move in the same direction termed ‘drift correlation’ and a stochastic component called ‘diffusive correlation’. These components suggest the mechanisms that coordinate movements, with drift correlation indicating external influences, and diffusive correlation pointing to social interactions. We use two case studies to highlight the ability of our approach both to quantify correlated movements in tracking data and to suggest the mechanisms that generate the correlation. First, we use an abrupt change in movement correlation to pinpoint the onset of spring migration in barren-ground caribou. Second, we show how spatial proximity mediates intermittently correlated movements among khulans in the Gobi desert. We conclude by discussing the linkages of our approach to the theory of collective motion. This article is part of the theme issue 'Collective movement ecology'.

The ongoing refugee crisis in Europe has seen many countries rush to construct border security fencing to divert or control the flow of people. This follows a trend of border fence construction across Eurasia during the post-9/11 era. This development has gone largely unnoticed by conservation biologists during an era in which, ironically, transboundary cooperation has emerged as a conservation paradigm. These fences represent a major threat to wildlife because they can cause mortality, obstruct access to seasonally important resources, and reduce effective population size.We summarise the extent of the issue and propose concrete mitigation measures.

The Mongolian Gobi-Eastern Steppe Ecosystem is one of the largest remaining natural drylands and home to a unique assemblage of migratory ungulates. Connectivity and integrity of this ecosystem are at risk if increasing human activities are not carefully planned and regulated. The Gobi part supports the largest remaining population of the Asiatic wild ass (Equus hemionus; locally called \"khulan\"). Individual khulan roam over areas of thousands of square kilometers and the scale of their movements is among the largest described for terrestrial mammals, making them particularly difficult to monitor. Although GPS satellite telemetry makes it possible to track animals in near-real time and remote sensing provides environmental data at the landscape scale, remotely collected data also harbors the risk of missing important abiotic or biotic environmental variables or life history events. We tested the potential of animal born camera systems (\"camera collars\") to improve our understanding of the drivers and limitations of khulan movements. Deployment of a camera collar on an adult khulan mare resulted in 7,881 images over a one-year period. Over half of the images showed other khulan and 1,630 images showed enough of the collared khulan to classify the behaviour of the animals seen into several main categories. These khulan images provided us with: i) new insights into important life history events and grouping dynamics, ii) allowed us to calculate time budgets for many more animals than the collared khulan alone, and iii) provided us with a training dataset for calibrating data from accelerometer and tilt sensors in the collar. The images also allowed to document khulan behaviour near infrastructure and to obtain a day-time encounter rate between a specific khulan with semi-nomadic herders and their livestock. Lastly, the images allowed us to ground truth the availability of water by: i) confirming waterpoints predicted from other analyses, ii) detecting new waterpoints, and iii) compare precipitation records for rain and snow from landscape scale climate products with those documented by the camera collar. We discuss the added value of deploying camera collars on a subset of animals in remote, highly variable ecosystems for research and conservation.

Most large herbivores in arid landscapes need to drink which constrains their movements and makes them vulnerable to disturbance. Asiatic wild ass or khulan (Equus hemionus) were widespread and abundant throughout the arid landscapes of Central Asia and Mongolia, but have undergone dramatic population declines and range constrictions; denying khulan access to water is believed to have played a major role. Mongolia’s South Gobi Region now houses the world largest remaining khulan population, but is undergoing rapid land use changes. Khulan water use is poorly understood, largely due to the difficulty of mapping waterpoints used by khulan throughout their exceptionally large ranges, prone to high variations in precipitation. We used the special movement path characteristics of GPS tagged khulan to show us where water is located. We identified 367 waterpoints, 53 of which were of population importance, characterized the seasonal and circadian use, and identified snow cover as the most important variable predicting khulan visits during the non-growing season, and vegetation greenness during the growing season. Our results provide a data layer to help guide a regional khulan conservation strategy, allow predictions for other part of the global khulan range, and illustrates the overall importance of waterpoints for dryland herbivores.

The microbial composition of host-associated microbiomes is influenced by co-evolutionary interactions, host genetics, domestication, and the environment. This study investigates the contribution of environmental microbiota from freshwater bodies to the gastrointestinal microbiomes of wild khulans ( Equus hemionus hemionus , n  = 21) and compares them with those of captive khulans ( n  = 12) and other equids—Przewalski’s horse ( n  = 82) and domestic horse ( n  = 26). Using PacBio technology and the LotuS pipeline for 16S rRNA gene sequencing, we analyze microbial diversity and conduct differential abundance, alpha, and beta diversity analyses. Results indicate limited microbial sharing between wild khulans and their waterhole environments, suggesting minimal environmental influence on their gut microbiomes and low levels of water contamination by khulans. Wild khulans exhibit greater microbial diversity and richness compared to captive ones, likely due to adaptations to the harsh nutritional conditions of the Gobi desert. Conversely, captive khulans show reduced microbial diversity, potentially affected by dietary changes during captivity. These findings highlight the significant impact of environment and lifestyle on the gut microbiomes of equids.

With increasing livestock numbers, competition and avoidance are increasingly shaping resource availability for wild ungulates. Shifts in the dietary niche of wild ungulates are likely and can be expected to negatively affect their fitness. The Mongolian Gobi constitutes the largest remaining refuge for several threatened ungulates, but unprecedentedly high livestock numbers are sparking growing concerns over rangeland health and impacts on threatened ungulates like the Asiatic wild ass (khulan). Previous stable isotope analysis of khulan tail hair from the Dzungarian Gobi suggested that they graze in summer but switch to a poorer mixed C3 grass / C4 shrub diet in winter, most likely in reaction to local herders and their livestock. Here we attempt to validate these findings with a different methodology, DNA metabarcoding. Further, we extend the scope of the original study to the South Gobi Region, where we expect higher proportions of low-quality browse in the khulan winter diet due to a higher human and livestock presence. Barcoding confirmed the assumptions behind the seasonal diet change observed in the Dzungarian Gobi isotope data, and new isotope analysis revealed a strong seasonal pattern and higher C4 plant intake in the South Gobi Region, in line with our expectations. However, DNA barcoding revealed C4 domination of winter diet was due to C4 grasses (rather than shrubs) for the South Gobi Region. Slight climatic differences result in regional shifts in the occurrence of C3 and C4 grasses and shrubs, which do not allow for an isotopic separation along the grazer-browser continuum over the entire Gobi. Our findings do not allow us to confirm human impacts upon dietary preferences in khulan as we lack seasonal samples from the South Gobi Region. However, these data provide novel insight into khulan diet, raise new questions about plant availability versuspreference, and provide a cautionary tale about indirect analysis methods if used in isolation or extrapolated to the landscape level. Good concordance between relative read abundance of C4 genera from barcoding and proportion of C4 plants from isotope analysis adds to a growing body of evidence that barcoding is a promising quantitative tool to understand resource partitioning in ungulates.

The Przewalski’s horse (Equus ferus przewalskii), the only remaining wild horse within the equid family, is one of only a handful of species worldwide that went extinct in the wild, was saved by captive breeding, and has been successfully returned to the wild. However, concerns remain that after multiple generations in captivity the ecology of the Przewalski’s horse and / or the ecological conditions in its former range have changed in a way compromising the species’ long term survival. We analyzed stable isotope chronologies from tail hair of pre-extinction and reintroduced Przewalski’s horses from the Dzungarian Gobi and detected a clear difference in the isotopic dietary composition. The direction of the dietary shift from being a mixed feeder in winter and a grazer in summer in the past, to a year-round grazer nowadays, is best explained by a release from human hunting pressure. A changed, positive societal attitude towards the species allows reintroduced Przewalski’s horses to utilize the scarce, grass-dominated pastures of the Gobi alongside local people and their livestock whereas their historic conspecifics were forced into less productive habitats dominated by browse.

Studying nomadic animal movement across species and ecosystems is essential for better understanding variability in nomadism. In arid environments, unpredictable changes in water and forage resources are known drivers of nomadic movements. Water resources vary temporally but are often spatially stationary, whereas foraging resources are often both temporally and spatially variable. These differences may lead to different types of nomadic movements: forage‐ vs. water‐driven nomadism. Our study investigates these two different types of nomadism in relation to resource gradients from mesic steppe to xeric desert environments in Mongolia's Gobi‐Steppe Ecosystem. We hypothesized that in the desert, where water is a key resource, animals are more water‐dependent and may show water‐driven nomadism with frequent revisits to spatially fixed resources, while in the steppe, animals are less water‐dependent and may show forage‐driven nomadism, tracking high‐quality foraging patches with infrequent revisits to previously used resources. We utilized GPS movement data from 40 individuals of four ungulate species (Mongolian gazelle, goitered gazelle, saiga antelope, and Asiatic wild ass) in the Gobi‐Steppe Ecosystem. We calculated displacement distances and recursion metrics and subsequently performed a principal component analysis to quantify the variation in movement patterns. The satellite‐derived vegetation greenness served as a proxy for the resource gradient and was associated with variation in movement behaviors described by the first principal component, demonstrating that the variability in movements was closely related to the resource gradient from mesic to xeric habitats. We showed that ungulates in the resource‐rich steppe tended to move long distances with few revisits (forage‐driven nomadism), while ungulates in the resource‐poor desert tended to move shorter distances with more revisits (water‐driven nomadism). Our results suggest that xeric and mesic habitats promote different types of nomadic strategies. These results have important implications for conservation strategies: Forage‐driven nomads primarily require a high degree of landscape‐level permeability, and water‐driven nomads additionally require the protection of ephemeral water bodies and actions to maintain the functional connectivity between them.