Tracking Five Millennia of Horse Management with Extensive Ancient Genome Time Series.

Horse domestication revolutionized warfare and accelerated travel, trade, and the geographic expansion of languages. Here, we present the largest DNA time series for a non-human organism to date, including genome-scale data from 149 ancient animals and 129 ancient genomes (≥1-fold coverage), 87 of which are new. This extensive dataset allows us to assess the modern legacy of past equestrian civilizations. We find that two extinct horse lineages existed during early domestication, one at the far western (Iberia) and the other at the far eastern range (Siberia) of Eurasia. None of these contributed significantly to modern diversity. We show that the influence of Persian-related horse lineages increased following the Islamic conquests in Europe and Asia. Multiple alleles associated with elite-racing, including at the MSTN "speed gene," only rose in popularity within the last millennium. Finally, the development of modern breeding impacted genetic diversity more dramatically than the previous millennia of human management.

Leaf trait plasticity reveals interactive effects of temporally disjunct grazing and warming on plant communities.

Changes in climate and grazing intensity influence plant-community compositions and their functional structure. Yet, little is known about their possible interactive effects when climate change mainly has consequences during the growing season and grazing occurs off growing season (dormant season grazing). We examined the contribution of trait plasticity to the immediate responses in the functional structure of plant community due to the interplay between these two temporally disjunct drivers. We conducted a field experiment in the northern Mongolian steppe, where climate was manipulated by open-top chambers (OTCs) for two growing seasons, increasing temperature and decreasing soil moisture (i.e., increased aridity), and grazing was excluded for one dormant season between these two growing seasons. We calculated the community-weighted mean (CWM) and the functional diversity (FD) of six leaf traits. Based on a variance partitioning approach, we evaluated how much of the responses in CWM and FD to OTCs and dormant season grazing occur through plasticity. The interactive effect of OTCs and the dormant season grazing were detected only after considering the role of trait plasticity. Overall, OTCs influenced the responses in CWM more than in FD, but the effects of OTCs were much less pronounced where dormant season grazing occurred. Thus, warming (together with decreased soil moisture) and the elimination of dormant season grazing could interact to impact the functional trait structure of plant communities through trait plasticity. Climate change effects should be considered in the context of altered land use, even if temporally disjunct.

Author Correction: Ancient hepatitis B viruses from the Bronze Age to the Medieval period.

In Fig. 2 of this Letter, the 'E' and 'G' clade labels were inadvertently reversed, and in Table 2 the genotype of DA27 was 'D1' instead of 'D5'. These have been corrected online.

Ancient hepatitis B viruses from the Bronze Age to the Medieval period.

Hepatitis B virus (HBV) is a major cause of human hepatitis. There is considerable uncertainty about the timescale of its evolution and its association with humans. Here we present 12 full or partial ancient HBV genomes that are between approximately 0.8 and 4.5 thousand years old. The ancient sequences group either within or in a sister relationship with extant human or other ape HBV clades. Generally, the genome properties follow those of modern HBV. The root of the HBV tree is projected to between 8.6 and 20.9 thousand years ago, and we estimate a substitution rate of 8.04 × 10-6-1.51 × 10-5 nucleotide substitutions per site per year. In several cases, the geographical locations of the ancient genotypes do not match present-day distributions. Genotypes that today are typical of Africa and Asia, and a subgenotype from India, are shown to have an early Eurasian presence. The geographical and temporal patterns that we observe in ancient and modern HBV genotypes are compatible with well-documented human migrations during the Bronze and Iron Ages1,2. We provide evidence for the creation of HBV genotype A via recombination, and for a long-term association of modern HBV genotypes with humans, including the discovery of a human genotype that is now extinct. These data expose a complexity of HBV evolution that is not evident when considering modern sequences alone.

Author Correction: 137 ancient human genomes from across the Eurasian steppes.

with In this Article, Angela M. Taravella and Melissa A. Wilson Sayres have been added to the author list (associated with: School of Life Sciences, Center for Evolution and Medicine, The Biodesign Institute, Arizona State University, Tempe, AZ, USA). The author list and Author Information section have been corrected online.

137 ancient human genomes from across the Eurasian steppes.

For thousands of years the Eurasian steppes have been a centre of human migrations and cultural change. Here we sequence the genomes of 137 ancient humans (about 1× average coverage), covering a period of 4,000 years, to understand the population history of the Eurasian steppes after the Bronze Age migrations. We find that the genetics of the Scythian groups that dominated the Eurasian steppes throughout the Iron Age were highly structured, with diverse origins comprising Late Bronze Age herders, European farmers and southern Siberian hunter-gatherers. Later, Scythians admixed with the eastern steppe nomads who formed the Xiongnu confederations, and moved westward in about the second or third century BC, forming the Hun traditions in the fourth-fifth century AD, and carrying with them plague that was basal to the Justinian plague. These nomads were further admixed with East Asian groups during several short-term khanates in the Medieval period. These historical events transformed the Eurasian steppes from being inhabited by Indo-European speakers of largely West Eurasian ancestry to the mostly Turkic-speaking groups of the present day, who are primarily of East Asian ancestry.

Metal-Tolerant Fungal Communities Are Delineated by High Zinc, Lead, and Copper Concentrations in Metalliferous Gobi Desert Soils.

The soil fungal ecology of the southern Gobi region of Mongolia has been little studied. We utilized the ITS1 region from soil DNA to study possible influences soil metal concentrations on soil fungal community variation. In the sample network, a distinctive fungal community was closely associated with high zinc (Zn), lead (Pb), and copper (Cu) concentrations. The pattern of occurrence suggests that high metal concentrations are natural and not a product of mining activities. The metal-associated fungal community differs little from the "normal" community in its major OTUs, and in terms of major fungal guilds and taxa, and its distinctiveness depends on a combination of many less common OTUs. The fungal community in the sites with high metal concentrations is no less diverse than that in areas with normal background levels. Overall, these findings raise interesting questions of the evolutionary origin and functional characteristics of this apparently "metal-tolerant" community, and of the associated soil biota in general. It is possible that rehabilitation of metal-contaminated mined soils from spoil heaps could benefit from the incorporation of fungi derived from these areas.

Antibody detection from Middendorf's vole (Microtus middendorffii) against Tula virus captured in Mongolia.

Mongolia in 2010 and 2011. A total of 76 voles belonging to the genera Myodes and Microtus were captured. Most of the voles that were seropositive to Tula virus antigen were Middendorf's voles (Microtus middendorffii (6/31)). Two of the 18 Myodes voles were also seropositive to Tula virus antigen. On the other hand, only one vole was seropositive to Puumala virus antigen. The results suggest that Tula virus was maintained in Middendorf's vole. This is the first report of detection of anti-Tula virus antibody in the central part of the Eurasia continent.

Soil and ecosystem respiration responses to grazing, watering and experimental warming chamber treatments across topographical gradients in northern Mongolia.

Globally, soil respiration is one of the largest fluxes of carbon to the atmosphere and is known to be sensitive to climate change, representing a potential positive feedback. We conducted a number of field experiments to study independent and combined impacts of topography, watering, grazing and climate manipulations on bare soil and vegetated soil (i.e., ecosystem) respiration in northern Mongolia, an area known to be highly vulnerable to climate change and overgrazing. Our results indicated that soil moisture is the most important driving factor for carbon fluxes in this semi-arid ecosystem, based on smaller carbon fluxes under drier conditions. Warmer conditions did not result in increased respiration. Although the system has local topographical gradients in terms of nutrient, moisture availability and plant species, soil respiration responses to OTC treatments were similar on the upper and lower slopes, implying that local heterogeneity may not be important for scaling up the results. In contrast, ecosystem respiration responses to OTCs differed between the upper and the lower slopes, implying that the response of vegetation to climate change may override microbial responses. Our results also showed that light grazing may actually enhance soil respiration while decreasing ecosystem respiration, and grazing impact may not depend on climate change. Overall, our results indicate that soil and ecosystem respiration in this semi-arid steppe are more sensitive to precipitation fluctuation and grazing pressure than to temperature change.

Leaf-trait plasticity and species vulnerability to climate change in a Mongolian steppe.

Climate change is expected to modify plant assemblages in ways that will have major consequences for ecosystem functions. How climate change will affect community composition will depend on how individual species respond, which is likely related to interspecific differences in functional traits. The extraordinary plasticity of some plant traits is typically neglected in assessing how climate change will affect different species. In the Mongolian steppe, we examined whether leaf functional traits under ambient conditions and whether plasticity in these traits under altered climate could explain climate-induced biomass responses in 12 co-occurring plant species. We experimentally created three probable climate change scenarios and used a model selection procedure to determine the set of baseline traits or plasticity values that best explained biomass response. Under all climate change scenarios, plasticity for at least one leaf trait correlated with change in species performance, while functional leaf-trait values in ambient conditions did not. We demonstrate that trait plasticity could play a critical role in vulnerability of species to a rapidly changing environment. Plasticity should be considered when examining how climate change will affect plant performance, species' niche spaces, and ecological processes that depend on plant community composition.

Climate change and grazing interact to alter flowering patterns in the Mongolian steppe.

Socio-economic changes threaten nomadic pastoralism across the world, changing traditional grazing patterns. Such land-use changes will co-occur with climate change, and while both are potentially important determinants of future ecosystem functioning, interactions between them remain poorly understood. We investigated the effects of grazing by large herbivores and climate manipulation using open-top chambers (OTCs) on flower number and flowering species richness in mountain steppe of northern Mongolia. In this region, sedentary pastoralism is replacing nomadic pastoralism, and temperature is predicted to increase. Grazing and OTCs interacted to affect forb flowering richness, which was reduced following grazing removal, and reduced by OTCs in grazed plots only. This interaction was directly linked to the soil moisture and temperature environments created by the experimental treatments: most species flowered when both soil moisture and temperature levels were high (i.e. in grazed plots without OTCs), while fewer species flowered when either temperature, or moisture, or both, were low. Removal of grazing increased the average number of graminoid flowers produced at peak flowering in Year 1, but otherwise grazing removal and OTCs did not affect community-level flower composition. Of four abundant graminoid species examined individually, three showed increased flower number with grazing removal, while one showed the reverse. Four abundant forb species showed no significant response to either treatment. Our results highlight how climate change effects on mountain steppe could be contingent on land-use, and that studies designed to understand ecosystem response to climate change should incorporate co-occurring drivers of change, such as altered grazing regimes.

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.

Plant response to climate change varies with topography, interactions with neighbors, and ecotype.

Predicting the future of any given species represents an unprecedented challenge in light of the many environmental and biological factors that affect organismal performance and that also interact with drivers of global change. In a three-year experiment set in the Mongolian steppe, we examined the response of the common grass Festuca lenensis to manipulated temperature and water while controlling for topographic variation, plant-plant interactions, and ecotypic differentiation. Plant survival and growth responses to a warmer, drier climate varied within the landscape. Response to simulated increased precipitation occurred only in the absence of neighbors, demonstrating that plant-plant interactions can supersede the effects of climate change. F. lenensis also showed evidence of local adaptation in populations that were only 300 m apart. Individuals from the steep and dry upper slope showed a higher stress/drought tolerance, whereas those from the more productive lower slope showed a higher biomass production and a greater ability to cope with competition. Moreover, the response of this species to increased precipitation was ecotype specific, with water addition benefiting only the least stress-tolerant ecotype from the lower slope origin. This multifaceted approach illustrates the importance of placing climate change experiments within a realistic ecological and evolutionary framework. Existing sources of variation impacting plant performance may buffer or obscure climate change effects.

Climate-driven marmot-plague dynamics in Mongolia and China.

The incidence of plague has rebounded in the Americas, Asia, and Africa alongside rapid globalization and climate change. Previous studies have shown local climate to have significant nonlinear effects on plague dynamics among rodent communities. We analyzed an 18-year database of plague, spanning 1998 to 2015, in the foci of Mongolia and China to trace the associations between marmot plague and climate factors. Our results suggested a density-dependent effect of precipitation and a geographic location-dependent effect of temperature on marmot plague. That is, a significantly positive relationship was evident between risk of plague and precipitation only when the marmot density exceeded a certain threshold. The geographical heterogeneity of the temperature effect and the contrasting slopes of influence for the Qinghai-Tibet Plateau (QTP) and other regions in the study (nQTP) were primarily related to diversity of climate and landscape types.

Conservation implications of elucidating the Korean wolf taxonomic ambiguity through whole-genome sequencing.

The taxonomic status of the now likely extirpated Korean Peninsula wolf has been extensively debated, with some arguing it represents an independent wolf lineage, Canis coreanus. To investigate the Korean wolf's genetic affiliations and taxonomic status, we sequenced and analysed the genomes of a Korean wolf dated to the beginning of the 20th century, and a captive wolf originally from the Pyongyang Central Zoo. Our results indicated that the Korean wolf bears similar genetic ancestry to other regional East Asian populations, therefore suggesting it is not a distinct taxonomic lineage. We identified regional patterns of wolf population structure and admixture in East Asia with potential conservation consequences in the Korean Peninsula and on a regional scale. We find that the Korean wolf has similar genomic diversity and inbreeding to other East Asian wolves. Finally, we show that, in contrast to the historical sample, the captive wolf is genetically more similar to wolves from the Tibetan Plateau; hence, Korean wolf conservation programmes might not benefit from the inclusion of this specimen.

Vulnerability of the northern Mongolian steppe to climate change: insights from flower production and phenology.

The semiarid, northern Mongolian steppe, which still supports pastoral nomads who have used the steppe for millennia, has experienced an average 1.7 degrees C temperature rise over the past 40 years. Continuing climate change is likely to affect flowering phenology and flower numbers with potentially important consequences for plant community composition, ecosystem services, and herder livelihoods. Over the growing seasons of 2009 and 2010, we examined flowering responses to climate manipulation using open-top passive warming chambers (OTCs) at two locations on a south-facing slope: one on the moister, cooler lower slope and the other on the drier, warmer upper slope, where a watering treatment was added in a factorial design with warming. Canonical analysis of principal coordinates (CAP) revealed that OTCs reduced flower production and delayed peak flowering in graminoids as a whole but only affected forbs on the upper slope, where peak flowering was also delayed. OTCs affected flowering phenology in seven of eight species, which were examined individually, either by altering the time of peak flowering and/or the onset and/or cessation of flowering, as revealed by survival analysis. In 2010, which was the drier year, OTCs reduced flower production in two grasses but increased production in an annual forb found only on the upper slope. The particular effects of OTCs on phenology, and whether they caused an extension or contraction of the flowering season, differed among species, and often depended on year, or slope, or watering treatment; however, a relatively strong pattern emerged for 2010 when four species showed a contraction of the flowering season in OTCs. Watering increased flower production in two species in 2010, but slope location more often affected flowering phenology than did watering. Our results show the importance of taking landscape-scale variation into account in climate change studies and also contrasted with those of several studies set in cold, but wetter systems, where warming often causes greater or accelerated flower production. In cold, water-limited systems like the Mongolian steppe, warming may reduce flower numbers or the length of the flowering season by adding to water stress more than it relieves cold stress.

Legumes mitigate ecological consequences of a topographic gradient in a northern Mongolian steppe.

Topography should create spatial variation in water and nutrients and play an especially important role in the ecology of water-limited systems. We use stable isotopes to discern how plants respond both to ecological gradients associated with elevation and to neighboring legumes on a south-facing slope in the semi-arid, historically grazed steppe of northern Mongolia. Out of three target species, Potentilla acaulis, Potentilla sericea, and Festuca lenensis, when >30 cm from a legume, all showed a decrease in leaf δ(15)N with increasing elevation. This, together with measures of soil δ(15)N, suggests greater N processing at the moister, more productive, lower elevation, and more N fixation at the upper elevation, where cover of legumes and lichens and plant-available nitrate were greater. Total soil N was greater at the lower elevation, but not lichen biomass or root colonization by AMF. Leaf δ(13)C values for P. acaulis and F. lenensis are consistent with increasing water stress with elevation; δ(13)C values indicated the greatest intrinsic water use efficiency for P. sericea, which is more abundant at the upper elevation. Nearby legumes (<10 cm) moderate the effect of elevation on leaf δ(15)N, confirming legumes' meaningful input of N, and affect leaf δ(13)C for two species, suggesting an influence on the efficiency of carbon fixation. Variation in leaf %N and %C as a function of elevation and proximity to a legume differs among species. Apparently, most N input is at upper elevations, pointing to the possible importance of grazers, in addition to hydrological processes, as transporters of N throughout this landscape.

Caddisflies (Trichoptera) of Mongolia: an updated checklist with faunistic and biogeographical notes.

To establish the biogeographic affinities of the caddisfly fauna of Mongolia, published records and results of our faunistic studies were analyzed. This study captured more than 47,000 adults collected from 386 locations beside lakes, ponds, streams/rivers, and springs in ten sub-basins of Mongolia using Malaise traps, aerial sweeping, and ultraviolet lights. In total, 201 species have been recorded, and approximately 269 species may occur in Mongolia according to our estimation. In a comparison of species richness for the family level, the Limnephilidae and Leptoceridae were the richest in species. The families Brachycentridae, Glossosomatidae, and Psychomyiidae had low species richness, but they included the most dominant species in terms of abundance and/or the percentage of occurrence in the samples from multiple sub-basins. Comparing the sub-basins, the Selenge had the highest Shannon diversity (H' = 3.3) and the Gobi sub-basin had the lowest (H' = 1.5). According to the Jaccard index of similarity, caddisfly species assemblages of Mongolia's ten sub-basins were divided into two main groups: One group includes the Selenge, Shishkhed, Bulgan, Tes, and Depression of Great Lakes sub-basins; the other group includes the Kherlen, Onon, Khalkh Gol, Valley of Lakes, and Gobi sub-basins. The majority of Mongolian species were composed of East Palearctic taxa, with a small percentage of West Palearctic and Nearctic representatives and an even smaller percentage from the Oriental region, suggesting that the Mongolian Gobi Desert is, and has been, a significant barrier to the distribution of caddisfly species between China and Mongolia.

Grazing and ecosystem service delivery in global drylands.

Grazing represents the most extensive use of land worldwide. Yet its impacts on ecosystem services remain uncertain because pervasive interactions between grazing pressure, climate, soil properties, and biodiversity may occur but have never been addressed simultaneously. Using a standardized survey at 98 sites across six continents, we show that interactions between grazing pressure, climate, soil, and biodiversity are critical to explain the delivery of fundamental ecosystem services across drylands worldwide. Increasing grazing pressure reduced ecosystem service delivery in warmer and species-poor drylands, whereas positive effects of grazing were observed in colder and species-rich areas. Considering interactions between grazing and local abiotic and biotic factors is key for understanding the fate of dryland ecosystems under climate change and increasing human pressure.

Alpha and beta diversity patterns of macro-moths reveal a breakpoint along a latitudinal gradient in Mongolia.

Little is known about the diversity and distribution patterns of moths along latitudinal gradients. We studied macro-moths in Mongolia along an 860 km latitudinal climatic gradient to gain knowledge on community composition, alpha, beta, and gamma diversity as well as underlying factors, which can be used as baseline information for further studies related to climate change. We identified 236 species of moths of ten families. Our study shows that the diversity of moths increased with the latitude, i.e., low species richness in the south and higher richness in the north. Moth community composition changed along the gradient, and we revealed a breakpoint of beta diversity that divided grassland and desert communities. In the desert, beta diversity was driven by species loss (i.e., nestedness), and few tolerant species existed with high abundance. In contrast, in the grassland, beta diversity was driven by species replacement with more unique species, (i.e., species which occurred only in one site). We found the lowest species diversity in the transitional zones dominated by few generalist species such as Agrotis ripae and Anarta trifolii. Low precipitation and an increasing number of grazing goats are drivers of species loss. We suggest different conservation strategies regarding the contrasting patterns of beta diversity in desert and grassland.