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Data collectionVirus-host data was collated from various sources. Major sources for the association databases included data shared by Olival et al4., Pandit et al.3, and Johnson et al.13. In data provided by Olival et al (assessed September 2019), host-virus associations have been assigned a score, based on detection methods and tests that are specific and more reliable. We used associations that have been identified as the most reliable (stringent data) from Olival et al4. In addition, a query in GenBank was run to parse out hosts reported for each GenBank submission for viruses presented in each of these three databases. Initially, for each virus name, taxonomic ID was identified using entrez.esearch function in biopython package. The taxonomic ID helped linked to the GenBank databases, identify the ICTV lineage and associated data in PubMed20,21. NCBI TaxID closely follows the ICTV database, but some recent changes in ICTV might not always be reflected in NCBI, so we manually checked names to ensure matching. This included virus genus and family information along with a standard virus name. Host data were aggregated based on the taxonomic ID and associated standard name. Finally, for each virus, a search was completed in PubMed to compile the number of hits related to the virus and their vertebrate hosts using the search terms below. The number of PubMed hits (PMH1) were used as a proxy for sampling bias3,13. The virus-host association data source is presented in supplementary code and data files (https://zenodo.org/record/5899054).$$ searchterm= (+virus_name+,[Title/Abstract])\ ANDleft(host,OR,hosts,OR,reservoir,OR,reservoirs,OR right.\ wild,OR,wildlife,OR,domestic,OR,animal,OR,animals,OR\ mammal,OR,bird,OR,birds,OR,aves,OR,avian,OR,avians\ left. OR,vertebrate,OR,vertebrates,OR,surveillance,OR,sylvaticright)$$Along with the PubMed terms we also queried the nucleotide database on PubMed using the taxonomic ID to find the number of GenBank entries for these viruses (PMH2). A correlation analysis between the PMH1 and PMH2 of well-recognized known viruses showed a high correlation with each other for us to safely use GenBank hits for novel viruses during the prediction stage of the model (Fig. S32).Development of ({{{{{{boldsymbol{G}}}}}}}_{{{{{{boldsymbol{c}}}}}}})
a. Centrality measures of observed network (({{{{{{boldsymbol{G}}}}}}}_{{{{{{boldsymbol{c}}}}}}}))To test if centrality measures (degree centrality, betweenness centrality, eigenvector centrality, clustering coefficient) for viral nodes in the observed network (({G}_{c})) vary significantly between viral families, we firstly used the Kolmogorov-Smirnov (KS) test. KS test is routinely used to identify distances between cumulative distribution functions of two probability distributions and is largely used to compare degree distributions of networks22,23. For each viral family, distributions of centrality measures (degree centrality, betweenness centrality, and eigenvector centrality) and clustering coefficient within the observed network (({G}_{c})) were compared with the distribution of all nodes in the network using the two-tailed KS test. Secondly, a linear regression model with virus family as a categorical variable and the number of PubMed hits as a covariate to adjust for sampling bias were fitted to understand associations of viral families with centrality measures.$${centrality},{measure}={beta }_{0}{intercept}+{{beta }_{1}{Viral}{family}}_{{categorical}}+{beta }_{2}{PubMed},{hits}$$After fitting the model, node-level permutations were implemented. For each random permutation, the output variable was randomly assigned to covariate values and the model was re-fitted. Finally, a p-value was calculated by comparing the distribution of coefficients from permutations with the original model coefficient.Network topology feature selectionUsing the observed network (({{{{{{boldsymbol{G}}}}}}}_{{{{{{boldsymbol{c}}}}}}})), multiple network topological features for all node (virus) pairs were calculated. The following are topological network features calculated. Features data type, definition and methods to calculate these features are presented in Table S3.1. The Jaccard coefficient: a commonly used similarity metric between nodes in information retrieval, is also called an intersection of over the union for two nodes in the network. In the unipartite network generated here, it represents the proportion of common neighbor viruses from the union of neighbor viruses for two nodes. Neighbor viruses are defined as viruses with which the virus shares at least a single host.2. Adamic/Adar (Frequency-Weighted Common Neighbors): Is the sum of inverse logarithmic degree centrality of the neighbors shared by two nodes in the network24. The concept of Adamic Adar index is a weighted common neighbors for viruses in the network. Within network prediction, the index assumes that viruses with large neighborhoods have a less significant impact while predicting a connection between two viruses compared with smaller neighborhoods.Both Jaccard and Adamic Adar coefficients have been routinely used for generalized network prediction and have shown high accuracy in predicting missing links in networks, specifically bipartite networks25, the information flowing through neighborhoods formed by two nodes might not always be enough to have similar predictive power in an unipartite network. This warrants use of other topology features along with neighborhood-based features.3. Resource allocation: Similarity score of two nodes defined by the weights of common neighbors of two nodes. Resource allocation is another measure to quantify the closeness of two nodes in the network and hence to understand the similarity of hosts they infect.4. Preferential attachment coefficients: The mechanism of preferential attachment can be used to generate evolving scale-free networks, where the probability that a new link is connected to node x is proportional to k26.5. Betweenness centrality: For a node in the network betweenness centrality is the sum of the fraction of all-pairs shortest paths that pass through it. The feature that we used for training the supervised learning model was the absolute difference between of betweenness centralities of two nodes. The difference between the betweenness centrality represents the difference in the sharing observed by two viruses in the pair.6. Degree centrality: The degree centrality for a node v is the fraction of nodes it is connected to. The feature that we used for training the supervised learning model was the absolute difference between degree centralities of two nodes. Unlike the difference in the betweenness centrality, the difference in degree centrality only looks at the difference in the number of observed host sharing.7. Network clustering: All nodes were classified into community clusters using Louvain methods27. A binary feature variable was generated to describe if both the nodes in the pair were part of the same cluster or not. If both viruses are from the same cluster, it represents a similar host predilection than when both viruses are not from the same cluster hence accounting for the evolutionary predilection of viruses (or virus families) to infect a certain type of host.These topological network characteristics come with certain limitations when it comes to the unipartite network of viruses with links formed due to shared hosts and might not truly represent the flow of information between nodes as compared to a bipartite network. Therefore, to account for these limitations, we use multiple network features as weak learners in our model building characteristics summarizing the network through the use of several quantitative metrics. In addition to this, we estimated the feature importance of these metrics in predicting missing links between viruses to quantify the information pasting through these links.Pearson’s correlation coefficients were calculated to identify highly correlated features and for choosing features for model training (Fig. S33). Virological features included in model training were categorical variables describing the virus family of both the nodes in the pair, followed by a binary variable if both the viruses belong to the same virus family. During the model development, PubMed hits generated three predictive features for each pair of viruses on which model training and predictions were conducted. These included two features representing PubMed hits for the two viruses in the pair (PubMedV1, PubMedV2) and the absolute difference between PubMedV1 and PubMedV2 to account for differences in sampling bias between the two viruses.Cross-validation and fitting generalized boosting machine (GBMs) modelsA nested-cross-validation was implemented for the binary model while simple cross-validation was implemented for the multiclass model (multiple output categories). The parameters of the binary model were first hyper-tuned using a cross-validated grid-search method. Values were tested using a grid search to find the best-performing model parameters that showed the highest sensitivity (recall). The parameters tested for hypertuning and their performance are provided in the supplementary material (supplementary results and Table S5). For further cross-validation of the overall binary model, all the viruses were randomly assigned to five groups. For each fold, the viruses assigned to a group were dropped from the data, and a temporary training network (({{{{{{boldsymbol{G}}}}}}}_{{{{{{boldsymbol{t}}}}}}}{{{{{boldsymbol{)}}}}}}) was constructed, assuming that this represented the current observed status of the virus-host community. For all possible pairs in ({{{{{{boldsymbol{G}}}}}}}_{{{{{{boldsymbol{t}}}}}}}) (both that sharing and not sharing any hosts) ten topological and viral characteristics were calculated as training features (Table S4). Categorical features were one-hot-encoded and numeric features were scaled. An XGBClassifier model with binary: logistic family was trained using the feature dataset to predict if virus pairs share hosts (1,0 encoded output). The cross-validation was also used to determine the optimum decision threshold for determining binary classification (Fig. S6) and a precision-recall curve was used to identify positive predictive value and sensitivity at the optimum threshold (Fig. S8).The multiclass model was implemented in the same way, creating an observed network (({G}_{c})) based on species-level sharing of hosts and randomly dropping viruses to generate a training network (({G}_{t})) to train the XGboost model. The output variables were generated based on the taxonomical orders of shared hosts. A pair of viruses can share multiple hosts, hence we trained a multioutput-multiclass model. Humans were considered an independent category of taxonomical order (label) and were given a separate label from primates. For fine-tuning the multiclass model, we started with the best performing parameters of the binary model and manually tested 5 combinations of model parameters by adjusting values of the learning rate, number of estimators, maximum depth, and minimum child weight (Supplementary code and results).We used three methods to estimate the importance of features for our binary model. Specifically, improvement in accuracy brought by branching based on the feature (gain), the percentage of times the feature appears in the XGboost tree model (weight), and the relative number of observations related to the specific feature (cover). Results for feature importance are shown in supplementary results (Fig. S10).Missing links for novel viruses, binary and multiclass predictionThe wildlife surveillance data represented a sampling of 99,379 animals (94,723 wildlife, 4656 domesticated animals) conducted in 34 countries around the world between 2009–2019 (Table S6)1. Specimens were tested using conventional Rt-PCR, Quantitative PCR, Sanger sequencing, and Next Generation Sequencing protocols to detect viruses from 28 virus families or taxonomic groups (Table S7). Testing resulted in 951 novel monophyletic clusters of virus sequences (referred to as novel viruses henceforth). Within 951 novel viruses, 944 novel viruses had vertebrate hosts that were identified with certainty based on barcoding methods and field identification. Host species identification was confirmed by cytochrome b (cytb) DNA barcoding using DNA extracted from the samples28. We predicted the shared host links between novel viruses and known viruses using binary and multiclass models in the following steps. Out of 944 novel viruses discovered in the last ten years, we were able to generate predictions for 531 novel viruses that were detected in species already classified as hosts within the network. The remaining 413 viruses were the first detection of any virus in that species and thus host associations could not be informed by the observed network (({{{{{{boldsymbol{G}}}}}}}_{{{{{{boldsymbol{C}}}}}}})) data.1. A new node representing the novel virus was inserted in the observed network (({{{{{{boldsymbol{G}}}}}}}_{{{{{{boldsymbol{c}}}}}}})). Using the list of species in which the novel virus was detected, new edges were created with known viruses that are also known to be found in those hosts. This generated a temporary network for the novel virus (({{{{{{boldsymbol{G}}}}}}}_{{temp}})). If the novel virus was not able to generate any edges with known viruses, meaning the host in which they have been found was never found positive for any known virus, predictions were not performed.2. Using ({{{{{{boldsymbol{G}}}}}}}_{{temp}}) feature values were calculated for the novel virus (betweenness centrality, clustering, and degree). For all possible pairs of the novel virus with known viruses that are not yet connected with each other through an edge in ({{{{{{boldsymbol{G}}}}}}}_{{temp}}) a feature dataset was generated (Jaccard coefficient(novel virus, known virus), the difference in betweenness centrality of the novel virus and known virus, if the novel virus and known virus were in the same cluster, the difference in degree centrality(novel virus, known virus), if the novel virus and known virus were from same virus family, the difference in PubMed hits(novel virus, known virus), PubMed hits for the novel virus, PubMed hits for the known virus). Studies and nucleotide sequences for novel viruses are expected to be published and shared on PubMed’s Nucleotide database and in various peer-reviewed publications. Data associated with GenBank accession numbers and nucleotide sequences for novel viruses are presented in Supplementary Data 3 and Supplementary Data 4 respectively. At the time of development of the model, data for all viruses was not shared in a format that would reflect on PubMed’s database, we decided to use the number of unique species the virus was detected in the last ten years of wildlife surveillance conducted by the USAID PREDICT project. These detections will be reflected in PubMed’s Nucleotide database and search term eventually, hence we considered them as a proxy for search terms conducted for known viruses. Currently, evaluation of the effects of this substitution of PubMed hits with the number of detections for novel viruses is not possible with limited data on novel viruses but needs to be reevaluated as more studies are published on these novel viruses. To further evaluate the association between PubMed hits through search term and Genbank hits, we ran a generalized linear regression model with PubMed hits as dependent variable and Genbank hits as intendent variable, accounting for virus families.$${{PubMed}}_{{Search}}left({log }right)={beta }_{0}{intercept}+{{beta }_{1}{Virus}{family}}_{{categorical}}+{beta }_{2}{Genbank},{hits},({log })$$The results indicated that Genbank hits had statistically significant predictive value in predicting PubMed hits (β = 0.72, p  More

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This study presents a comprehensive quantification of carbon sequestration as well as CO2/CH4/N2O emissions reductions from terrestrial ecosystems based on multiple sources of data from literature, inventories, public databases and documents. The pathways considered ecosystem restoration and protection from being converted into cropland or built-up areas, reforestation, management with improved nitrogen use in cropland, restricted deforestation, grassland recovery, reducing risk from forest wildfire and others. Here we describe the cross-cutting methods that apply across all 16 NCS pathways. The definitions, detailed methods and data sources for evaluating individual pathways can be found in the Supplementary Information.Cross-cutting methodsBaseline settingWe set 2000 as the base year because the large-scale national ecological projects, such as the Grain for Green Project, were started since then. We first evaluate the historical mitigation capacity during 2000–2020, which is the first 20 years of implementing the projects. From this procedure we can determine how much mitigation capacity has been realized through the previous projects in the past two decades and to what extent additional actions can be made after 2020. Relative to the baseline 2000–2020, we then evaluate the maximum potentials of the NCS mitigation in the future 10 (2020–2030) and 40 (2020–2060) years, corresponding to the timetable of China’s NDCs: carbon peak before 2030 and carbon neutrality by 2060.The settings of baseline in this study are different from the existing assessments (2000s–2010s as a baseline and 2010–2025/2030/2050 as scenarios)1,22,23,27,28. Baseline sets the temporal and spatial reference for NCS pathway scenarios, which may have a great impact on the NCS estimates. Notably, NCS actions during 2000–2020 will have a great impact in the future periods, which we refer to as the ‘legacy effect’. The legacy effect itself, mainly reforestation, is independent of being assessed, but it is conceptually attributed to natural flux and excluded from future NCS potential estimates.Maximum potentialThe MAMP refers to the additional CO2 sequestration or avoided GHG emissions measured in CO2 equivalents (CO2e) at given flux rates in a period on the maximum extent to which the stewardship options are applied (numbers are expressed as TgCO2e yr−1 for individual pathways and PgCO2e yr−1 for national total) (Extended Data Fig. 1 and Supplementary Table 2). ‘Additional’ means mitigation outcomes due to human actions taken beyond business-as-usual land-use activities (since 2020) and excluding existing land fluxes not attributed to direct human activities1. The MAMP of CH4 and N2O are accounted by three cropland and wetland pathways (cropland nutrient management, improved rice cultivation and peatland restoration). We adopt 100 yr global warming potential to calculate the warming equivalent for CH4 (25) and N2O (298), respectively38,39 because these values are used in national GHG inventories, although some researchers have argued that using the fixed 100 yr global warming potential to calculate the warming equivalents may be problematic because they cannot differentiate the contrasting impacts of the long- and short-lived climate pollutants39. Because the flux rate of the GHG by ecosystems may vary with the time of recovery or growth, the MAMP may also change for different periods even given the same extent.The ‘maximum’ is constrained by varied factors across the NCS pathways. We constrain forest and grassland restoration by the rate of implementation, farmland red line and tree surviving rate (Extended Data Fig. 2). Surviving rate here is the ratio of the area with increased vegetation cover due to reforestation to the total reforestation area. The farmland red line refers to ‘the minimum area of cultivated land’ given by the Ministry of Land and Resources of China. It defines the lowest limit, and the current red line is ~120 Mha. It is a rigid constraint below which the total amount of cultivated land cannot be reduced. From this total amount, there is provincial farmland red line. This red line sets a constraint on the implementation of the NCS pathways associated with land-use change. We set the future scenario of farmland area that can be used for grassland or forest restoration on the basis of the provincial farmland red line. Basic farmland is closely related to national food security. By 2050, China’s population is predicted to decrease slightly, but with economic development, the per capita demand for food may increase40. We assume that the food production in the future can meet the food demand via increasing agricultural investment and technological advancement. The N fertilizer reduction scenario is set to be below the level 60%, under which crop yield is not significantly affected19, because N fertilizer is surplus in many Chinese croplands. For timber production, we assume that the demand for timber can be met if the production level is maintained at the level of 2010–2020 (83.31 million m3 yr−1). As deforestation of natural forests is 100% forbidden since 2020, the future timber will come mainly from tree plantations. For grazing optimization, we assume that livestock production is not affected by grassland fencing due to refined livestock management such as improving feed nutrient and fine-seed breeding41.The areas of historical NCS implementation during 2000–2020 were estimated using statistical data, published literature and public documents, with a supplement from remote-sensing data. The flux rates were obtained either by directly using the values from multiple literature sources or from estimates using the empirical formulae. For the estimates of future NCS potential, the flux rate and extent of the pathway were determined on the basis of the baseline (2000–2020). The extent is assumed to be achieved by using the same rate but limited by the multiple constraints stated in the preceding unless the implementation scopes have been reported in national planning documents. We estimate the legacy effect by multiplying the implementation area in the past by the flux rates in the future two periods.SaturationThe future mitigation potential that we estimate for 2030 and 2060 will not persist indefinitely because the finite potential for natural ecosystems to store additional carbon will saturate. For each NCS pathway, we estimate the expected duration of the potential for sequestration at the maximum rate (Supplementary Table 3). Forests can continue to sequester carbon for 70–100 years or more. Restored grasslands and fenced grasslands can continue to sequester carbon for >50 years. Forest-fire management and cover crops can continue to sequester carbon for 40–50 years or more. Sea grasses and peatlands can continue to sequester carbon for millennia. Avoided pathways do not saturate as long as the business-as-usual cases indicate that there are potential areas for avoided losses of ecosystems. In this case, sea grass and salt marsh would disappear entirely after 64 years, but it would be 100–300 years or more for forest, grassland and peatland.Estimation of uncertaintiesThe extent (area or biomass amount) and flux (sequestration or reduced emission per area or biomass amount in unit time) are considered to estimate uncertainty of the historical mitigation capacity or future potential for each NCS pathway. We use the IPCC approaches to combine uncertainty42. Where mean and standard deviation can be estimated from collected literature, 95% CIs are presented on the basis of multiple published estimates. Where a sample of estimates is not available but only a range of a factor, we report uncertainty as a range and use Monte Carlo simulations (with normal distribution and 100,000 iterations) to combine the uncertainties of extent and flux (IPCC Approach 2). The overall uncertainties of the 16 NCS pathways were combined using IPCC Approach 142. If the extent estimate is based on a policy determination, rather than an empirical estimate of biophysical potential, we do not consider it a source of uncertainty.MACsThe economic/cost constraints refer to the amount of NCS that can be achieved at a given social cost. The MAC curve is fitted according to the total publicly funded investment and total mitigation capacity or potential during a period. The MAC curves are drawn to estimate the historical mitigation or MAMP at the cost thresholds of US$10, US$50 and US$100 (MgCO2e)−1, respectively. The trading price in China’s current carbon market is ~US$10 USD (as the minimum cost43), and the cost-effective price point44,45 to achieve the Paris Agreement goal of limiting global warming to below 2 °C above pre-industrial levels is US$100 (as the maximum cost). A carbon price of US$50 is regarded as a medium value1,46. For the pathways of reforestation, avoided grassland conversion, grazing optimization and grassland restoration, we collected the statistical data of investments in China from 2000 to 2020 and estimated the affordable MAMP below the three mitigation costs. Due to data limitations, the points used for fitting the MAC curve are values for cost (invested funds) and benefit (mitigation capacity) in each of the provinces. We rank the ratio of benefit to cost in a descending order to obtain the maximum marginal benefit for MAC by assuming that NCS measures are first implemented in the region with the highest cost/benefit rate. We refer to the investment standard before 2020 as the benchmark and estimate the cost of each pathway for the future periods with discount rates of 3% and 5%, respectively. The social discount rate 4–6% is usually used as a benchmark discount value in carbon price studies in China compared with lower scenarios (for example, 3.6%)46,47. In a global study for estimating country-level social cost of carbon, 3% and 5% are used for scenario analysis48. Note that the mean value from the two discount rates was used in presenting the results. For the other pathways where investment data cannot be obtained, we refer to relevant references to estimate MAC. All the cost estimates are expressed in 2015 dollars, transformed on the basis of the Renminbi and US dollar exchange rate of the same year. The year 2015 represents a relatively stable condition of economic increase over the past decade (2011–2020) in China (the increase rate of gross domestic product (GDP) in 2015 is similar to the 10 yr mean). In the cases when the MAC curves exceed the estimated maximum potentials in the period, we identify the historical capacity or the MAMP as limited by the biophysical estimates.Additional mitigation required to meet Paris Agreement NDCsOn 28 October 2021, China officially submitted ‘China’s Achievements, New Goals and New Measures for Nationally Determined Contributions’ (‘New Measures 2021’ hereafter) and ‘China’s Mid-Century Long-Term Low Greenhouse Gas Emission Development Strategy’ to the Secretariat of the United Nations Framework Convention on Climate Change as an enhanced strategy to China’s updated NDCs (first submission in 2015). The goal of China’s updated NDCs is to strive to peak CO2 emissions before 2030 and achieve carbon neutralization by 2060. It specified the goals to include the following: before 2030, China’s carbon dioxide emissions per unit of GDP are expected be more than 65% lower than that in 2005, and the forest stock volume is expected to be increased by around 6.0 (previously 4.5) billion m3 over the 2005 level. In the ‘New Measures 2021’9 and ‘Master Plan of Major Projects of National Important Ecosystem Protection and Restoration (2021–2035)’5, many NCS-related opportunities are proposed to consolidate the carbon sequestration of ecosystems and increase the future NCS potential, including protecting the existing ecosystems, implementing engineering to precisely improve forest quality, continuously increasing forest area and stock volume, strengthening grassland protection and recovery and wetland protection and improving the quality of cultivated land and the agricultural carbon sinks.Industrial CO2 emissionsThe historical CO2 emissions data from 2000 to 201749,50 are used as the benchmark of industrial CO2 emissions during 2000–2020. For future projections, we use the peak value of the A1B2C2 scenario (in the range of 10,000 to 12,000 Mt) in 2030 from ref. 11. We assume that CO2 emission increases linearly from 2017 to 2030.Characterizing co-benefitsNCS activities proposed in the future measures or plans may enhance co-benefits. Four generalized types of ecosystem services are identified: improving biodiversity, water-related, soil-related and air-related ecosystem services (Fig. 1). Biodiversity benefits refer to the increase in different levels of diversity (alpha, beta and/or gamma diversity)51. Water, soil and air benefits refer to flood regulation and water purification, improved fertility and erosion prevention, and improvements in air quality, respectively, as defined in the Millennium Ecosystem Assessment52. The evidence that each pathway produces co-benefits from one or more peer-reviewed publications was collected through reviewing the literature (see the details for co-benefits of each pathway in Supplementary Information).Mapping province-level mitigationThe data for extent of implementing forest pathways are obtained from the statistical yearbook and reported at the province level. To be consistent with the forest pathways, the other pathways were also aggregated to the provincial-level estimate from the spatial data. If the flux data were available in different climate regions, the provinces are first assigned to climate regions. When a province spans multiple climate zones, the weight value is set according to the proportion of area, and finally an estimated value of rate was calculated (for fire management, some grassland and wetland pathways). For the forest pathways, we first collected the flux-rate data from reviewing literature and then averaged these flux rates to region/province. The flux rates for reforestation and natural forest management were calculated separately by province and age group. Similarly, specified flux rates are applied for different times after ecosystem restoration or conversion for other pathways.Classification of NCS typesThree types of NCS pathways were classified: protection (of intact natural ecosystems), improved management (on managed lands) and restoration (of native cover)35. In our study, four (AVFC, AVGC, AVCI, AVPI), eight (IMP, NFM, FM, BIOC, CVCR, CRNM, IMRC, GROP) and four (RF, GRR, CWR, PTR) NCS pathways were identified as protection, management and restoration types, respectively (Supplementary Table 1). These pathways can be further divided into groups of ‘single’ type or ‘mixed’ type according to their contribution to individual pathways. Specifically, in a certain area, when the mitigation capacity of a certain pathway accounts for more than 50% of the total, it is regarded as a single or dominant NCS type; if no single pathway accounts for more than 50%, it is a mixed type, named by the top pathways whose NCS sum exceeds 50% of the total mitigation capacity. More

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