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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

Influence of independent variables on the extent of stereotyped behaviourThe overall level of stereotypy we observed was low, suggesting that the pandas in our study were not seriously stressed. The variables that we found to be correlated with stereotypy are consistent with what we know of pandas’ natural history. Our study reports that variables like logs on the ground, nest, sociality, zoo, tree density, age and tree height used by pandas are the driving force for stereotypy in captive pandas involved in the study.Making the captive environment more naturalistic by integrating enrichment into the enclosure seems to be a promising way of alleviating stress and improving both welfare and reintroduction success41. It also helps to improve reproductive rate and overall health39. Improved health reduces stress and gives greater control over the environment increasing the chances of survival and longevity both in captivity and following release into the wild5. It is generally accepted that enrichment of the captive environment increases animals’ ability to cope with challenges and positive use of the environment reduces or eliminates aberrant behaviour23. Lack of enclosure enrichments and less complex enclosures can cause stereotypy and other atypical behaviours24, while providing enrichment increases the frequency of natural behaviours25 and thereby reduces stress, which in turn decreases stereotypy27. But enrichment needs to be appropriate for the species of animal concerned. Abnormal behaviours are often associated with captive conditions that deviate greatly from the species’ natural environment. Consistent with this argument we found that though dead and fallen logs on the ground are one of the important characteristics of the panda habitats in the wild42,43,44,45, merely providing them in captivity does not ensure the species’ welfare: in fact, stereotypy increased with log density in our study subjects. This could be due to the fact that four individuals that showed more stereotypy were housed in the small barren enclosures with no trees but more logs as a part of enrichment. Without those four individuals, the linear relation between stereotypy and log density was not statistically significant. This clearly suggested that merely providing logs in the small enclosures does not maintain welfare.
When animals are housed in enclosures designed to resemble their natural habitat by considering their natural history (provision of vegetation, shelter, pool, etc.), there is a reduction or elimination of abnormal patterns of behaviour such as stereotypies, increased fitness and improved health, all of which may influence reproduction25,46,47,48. For many species, nests, shelter or burrows in enclosures will serve as retreat and hiding places, which are essential to cope with environmental stressors10. Gerbils, mice and rabbits have all shown less stereotyped behaviour when retreats are provided9,49,50,51. Such retreats can mitigate the effects of zoo visitors, who can serve as a source of stress for species that rarely interact with humans in the wild. Consistent with these previous results, we found that with provision of nests, the extent of stereotypy decreased in captive pandas. Many species prefer nests both for rearing the young as well as for resting and shelter, and pandas follow this pattern, so providing nests in adequate numbers will supports their natural behaviour as well as provide relief from environmental stressors. Zidar recommends providing one more nest than there are individuals in an enclosure52.Although pandas are an asocial species, our study showed that pandas show more stereotyped behaviour when housed alone than when with another individual or in group. Being a solitary species in the wild might encourage management to house them singly in captivity, but not every activity and habit of species in the wild can be used in captivity. For example, polar bears are also a solitary species, and it was at one time thought best to manage them alone, but it was found that managing them in a social setting reduces stereotypic pacing behaviour53, consistent with this study. Importantly, managers of zoo should note that living in group is greatly influenced by the individuals’ compatibility and hence this should be kept in mind while pairing.Similarly, we found that the presence of trees, and greater mean tree height use by pandas, reduced stereotypy. Pandas’ preferred high elevation habitat is favourable for taller trees20, and Shrestha et al. found that canopy cover was an important factor in habitats for pandas in the wild54. In European zoos, pandas spend 90% of their time off the ground37. Consistent with these previous findings, our study reveals that more and taller trees support natural behaviours in panda. The Central Zoo Authority (CZA) of India enrichment manual recommends taller tree provision in panda enclosures, and again we provide empirical support for its recommendation.We found that with increasing age stereotypy increased in pandas. The older the individuals the more time spent in captivity with its associated risks of stereotypic behaviour. The same trend has been observed in other species: for example in captive bears stereotypic behaviour increased with age55. In another study Asiatic black bear and sun bear showed more stereotypy with age56.Influence of independent variables on behavioural diversityAs noted in the “Introduction” section, in a species like the panda, high daytime behavioural diversity is not necessarily a positive indication of good welfare. However, our comparison of behavioural diversity with stereotypy showed a negative trend (though not significant), suggesting that low behavioural diversity might be associated with poorer welfare.Nonetheless, we found some results that suggested that lower diversity might in fact be associated with a more natural lifestyle. Because of the amount of time that wild pandas spend foraging57 and sleeping or inactive, they cannot show much behavioural diversity, and in our sample of captive individuals, they showed the same trend. For example, behavioural diversity was lower when pandas were provided with more trees in the enclosure. This suggests that when appropriate conditions are maintained in captivity, panda prefer to be inactive during the day, as is consistent with their natural history57. As pandas are essentially arboreal mammals, naturally they also spend most of the time inactive (e.g. sleeping) on the trees57. Indeed, providing larger trees would promote inactive behaviours and hence lower behaviour diversity in captivity, this captures their natural behaviour. This is consistent with our results where increased tree height used by pandas decreased behavioural diversity.We found behavioural diversity was greater when there are more logs in the enclosure. In the Yele Reserve in Sichuan, China, Wei et al. found 107 of 185 panda dropping sites (57.8%) on shrub branches, 49 (26.5%) on fallen logs, and only 29 (15.7%) on the forest floor44. Droppings were found mostly on elevated structures ranging from 1 to 3 m above the forest floor and occasionally on trees over 12 m. Moreover, microhabitats selected by pandas were also characterized by fallen logs and tree stumps42,45. Wei and Zhang mention that to access bamboo leaves easily, pandas usually use some elevated objects, such as shrub branches, fallen logs, or tree stumps to lift their body43. Hence, providing tree logs in the vicinity supports their natural behaviour. But at the same time management should keep in mind that merely providing logs in the enclosure would not guarantee species welfare, as discussed in previous section with respect to stereotypy.Temperature is an important element of microclimate for animals, and influences the activity level of captive animals10. When temperature rises, many species show distress in captivity10. The red panda inhabits low-temperature areas20, so it is unlikely that higher temperatures would support natural behaviours. We found that with increased temperature behavioural diversity decreased in captive pandas. Similarly, we found that pandas showed higher behavioural diversity in the winter season, where temperatures are low as compared to summer season.Studies that have tried to relate behavioural diversity and stereotypy in captive animals have varied in their interpretation; many have found significantly inverse relationships between the two19. In this study our multivariate model suggested that behavioural diversity is negatively influenced by stereotypy in captive pandas, confirming previous research.Other factors associated with variations in behavioural diversity are less easy to identify with welfare, positive or negative. Behavioural diversity also decreases with age of pandas and increases with distance to cage mate, number of visitors and quantum of bamboo provided, though these effects were not significant in the REVS model.Taken together, these results suggest that higher behavioural diversity is not a straightforward indicator of better welfare in all captive animals. The overall non-significant relationship between stereotyped behaviour and diversity we observed could well be the result of a mixture of factors operating in opposite directions. To interpret diversity correctly, it would be helpful to know what level of diversity the species shows in the wild, and such data are rarely available—a limitation of our study as of many others. Although there are dissenting voices58, arguably what matters most both in terms of welfare and in terms of potential reintroduction to the wild, is that a captive animal’s time budget approximates as closely as possible that of a wild animal. It is not diversity as such that is important, but the behaviours that the animal exhibits.Differences between zoosOur study showed that both the extent of stereotyped behaviour and behavioural diversity varied significantly among zoos. However, Zoo 2, an important breeding centre, housed only a female and her two cubs; this may lead to many factors being confounded and is thus a limitation to our study. Captive animals rely on the zoo environment, its routine and husbandry practices to limit their stress levels, and any failure to provide suitable resources will certainly disturb them and lead to distress10. Controlling such variables appropriately will help reduce stress among captive animals, and we can rely to some extent on our knowledge of the species’ natural history to guide us through this challenge. Our study was able to identify some of the factors that are associated with better welfare, but even with these factors taken into account, significant differences among the three zoos remained. These are presumably due to subtler variations in the zoos’ environment or management regimes. Since the panda is endemic to high elevations, we considered whether differences between the elevations of the zoos might be relevant, but the biggest differences were between Zoos 1 and 3, which are at essentially the same elevation.In Zoo 1 pandas showed lower stereotypy and higher behavioural diversity then the other two zoos. Again, these differences may be due to subtle differences between the management regimes in the three zoos; possibilities include keepers’ attitudes and the zoo’s experience in managing pandas. It is notable that Zoo 1 has longer and wider experience in the management of red pandas than the other two zoos, which have joined the captive breeding programme more recently and have fewer animals. Other notable differences were that in Zoo 1, pandas are fed twice a day as compared to the other two zoos where feed is given all at one time (both bamboo and supplementary diet); and that in Zoo 1 the enclosures were of a good size for a small mammal like the red panda, and were well maintained with much natural vegetation. The other two zoos had a large enclosure with poor vegetation (trees and grass), or a small enclosure with a barren floor and no trees at all. Location of the enclosure also needs to be considered: in two of the enclosures at Zoo 3 the sun shone directly on the animals with no shade as such, keeping the temperature higher than would be natural for pandas. Any of these factors could be the reason the pandas performed comparatively well in Zoo 1, and it would be necessary to study a wider (and, therefore, cross-national) sample of zoos holding pandas to identify which of them are the most important. More

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