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    Network motifs shape distinct functioning of Earth’s moisture recycling hubs

    UTrack atmospheric moisture tracking modelThe UTrack atmospheric moisture tracking model is a novel Lagrangian model that tracks parcels of moisture forward in three-dimensional space9. UTrack is the first moisture tracking model to employ ERA5 reanalysis data8. The basic principle of the model is that for each mm of evaporation, a certain number of “moisture parcels” is released and subsequently tracked through time and space. At each time step, the moisture budget of the parcels is updated based on evaporation and precipitation at the respective time and location, meaning that for each location of evaporation, a detailed image of the “footprint” of evaporation can be created. All types of evapotranspiration are included, and here is simply called evaporation.For each mm of evaporation, 100 parcels are released 50 hPa above the surface height at random spatial locations within each 0.25° grid cell of input evaporation data. The trajectories of the parcels are based on interpolated three-dimensional ERA5 wind speed and wind direction data, which also have a horizontal resolution of 0.25° and consist of 25 pressure layers in the atmospheric column. The spatial coordinates of each parcel are updated at each time step of 0.1 h. Also, at each time step, there is a certain probability that a parcel is redistributed randomly along the atmospheric column such that, on average, every parcel is redistributed every 24 h (see methods section Moisture recycling dataset: validation and uncertainties below for further details). The relative probability of the new position in the atmospheric column is scaled with the vertical moisture profile. Parcels are tracked for 30 days or until 99% of their moisture has precipitated.To allocate a certain fraction of any moisture parcel to precipitation events at the current time and location, ERA5 hourly total precipitation (P) and total precipitable water (TPW) are interpolated to the simulation time step of 0.1 h. The amount of moisture that precipitates at a certain time step equals the amount of precipitation at that time step over the total precipitable water in the atmospheric water column (P/TPW). Specifically, precipitation A in mm per time step at location x, y at time t that originated as evaporation from a particular source is described as:$${A}_{x,y,t}={P}_{x,y,t}frac{{W}_{{{{{{{{rm{parcel,t}}}}}}}}}{E}_{{{{{{{{rm{source,t}}}}}}}}}}{{{{{{mathrm{TP}}}}}}{{{{{{mathrm{W}}}}}}}_{x,y,t}}$$
    (1)
    with P being precipitation in mm at time step t, Wparcel,t (mm) the amount of moisture in the parcel of interest, Esource,t the fraction of moisture present in the parcel at time t that has evaporated from the source, and TPW (mm) the precipitable water in the atmospheric water column. The moisture content of parcels is updated each time step using evaporation and precipitation at its current location:$${W}_{{{{{{{{rm{parcel,t}}}}}}}}}={W}_{{{{{{{{rm{parcel,t-1}}}}}}}}}+({E}_{{{{{{{{rm{x,y,t}}}}}}}}}-{P}_{{{{{{{{rm{x,y,t}}}}}}}}})frac{{W}_{{{{{{{{rm{parcel,t-1}}}}}}}}}}{{{{{{mathrm{TP}}}}}}{{{{{{mathrm{W}}}}}}}_{{{{{{{{rm{x,y,t}}}}}}}}}}$$
    (2)
    The moisture (fraction) that has evaporated from the source is updated as follows:$${E}_{{{{{{{{rm{source,t}}}}}}}}}=frac{{E}_{{{{{{{{rm{source,t-1}}}}}}}}}{W}_{{{{{{{{rm{parcel,t-1}}}}}}}}}{A}_{x,y,t}}{{W}_{{{{{{{{rm{parcel,t}}}}}}}}}}$$
    (3)
    The moisture flow mij from evaporation in cell i to precipitation in cell j is aggregated on a monthly basis (mm/month), where [x, y] ∈ j becomes:$${m}_{ij}=mathop{sum }limits_{t=0}^{{{{{{{{rm{month}}}}}}}}}{A}_{j,t}frac{{E}_{i,t}}{{W}_{i,t}}$$
    (4)
    with Wi,t being the tracked amount of moisture from the source cell i at time t. These simulations were performed for all evaporation on Earth during 2008–2017. The results were then aggregated on a mean-monthly basis to produce monthly means, and stored at 0.5 degree resolution. This dataset can be downloaded from ref. 53. For details on how to process the data, we refer to the accompanying paper by ref. 3.Moisture recycling dataset: validation and uncertaintiesAs with all moisture recycling simulations, the ones used in this study rely on a number of assumptions that may affect the moisture recycling rates. All offline moisture recycling models use atmospheric model output to simulate the path of evaporation through the atmosphere to the location where it precipitates. Therefore, there are two sources of uncertainty that affect the moisture recycling estimates: (1) the quality of the atmospheric forcing data and (2) the assumptions in the moisture tracking model.Tuinenburg and Staal (2020)9 explored these sources of uncertainty for a number of locations globally. The effects of a decrease in the quality of the atmospheric forcing data were most important in the vertical resolution of the atmospheric data: the forcing data should have enough vertical levels to resolve any vertical shear in atmospheric moisture transport. If the forcing data has a low vertical resolution, the moisture tracking model is forced with the mean atmospheric flow over a number of layers. In many regions, there are surface moisture flows that are in a different direction than the moisture flow aloft, resulting in a very small vertically integrated transport, which would distort the simulation of atmospheric moisture transport. Compared to the vertical resolution of the forcing data, the horizontal and temporal resolutions were less important in order to keep errors as small as possible. Because of the importance of this high vertical resolution, it was recommended9 to use the ERA5 dataset8 as its forcing dataset, as this currently is the atmospheric reanalysis dataset with the highest vertical resolution.In addition, the change of ERA-interim to ERA5 resulted in a much better land-surface scheme with monthly varying vegetation and better bare soil evaporation. Also, many more observations are assimilated, which results in a better precipitation product compared to ERA-interim. Following this, the tracking of atmospheric moisture using ERA5 allows for a better quality of the atmospheric moisture cycle than before. But, of course, also the already high horizontal resolution of 0.5∘ × 0. 5∘ has the limitation that very localized moisture recycling features like orography and locally varying land use cannot be resolved. Out of these reasons, the uncertainty in the evaporation estimates is a lot larger than that in the precipitation estimates, because of the lack of global evaporation measurements and the difficulty in measuring evaporation in general54,55.There are also uncertainties due to the assumptions in the moisture tracking model that can be split into a category of simulation assumptions and physical assumptions. The simulation assumptions include model formulation (Eulerian vs. Lagrangian model set-ups), time step lengths, number of parcels released, and types of interpolation. Of these simulation assumptions, the most important aspects were the model formulation, with Lagrangian models better able to resolve complex terrain and atmospheric flows. For the other model assumptions (see methods section UTrack atmospheric moisture tracking model), it was chosen to simulate with the highest level of precision before any more information (e.g., more parcels) would no longer affect evaporation footprints and moisture recycling statistics (see ref. 9 for further details). Even though the ERA5 dataset is known to have some precipitation biases in the tropics, the results of UTrack (forced by ERA5) have recently been validated across the tropics by independent measurements of deuterium excess, a measure of a stable isotope that depends on terrestrial precipitation recycling56. UTrack estimates and isotope-based estimates of terrestrial moisture recycling corresponded, especially in tropical rainforests (Kendall’s (overline{tau }=0.52)56), which are found to be moisture recycling hubs on a global scale.Network constructionMotivated by the network-like structure of the data, we here employ a network perspective to study moisture flows. Hence, nodes in such a network are grid cells on a regular spherical grid and edges represent the moisture transported. However, interpreting the dataset directly as a weighted network is neither computationally feasible nor does a weighted network allow for identifying motifs, the building blocks of complex networks17. We, therefore, aim for an approach utilizing an unweighted network.As shown in Fig. S1, moisture recycling strengths are heterogeneously distributed over multiple powers of magnitude. Thus, it is not appropriate to just withdraw the moisture transport volume and include all moisture transport connections within the dataset as equal and unweighted links. Instead, we attempt to highlight the strongest moisture pathways and, thus, the backbone of the Earth’s moisture recycling network. To, on the one hand, include as much moisture volume as possible but also keep the absolute volume of moisture transport represented per edge as similar as possible, we decided to include edges in a data-adaptive way: we step-wise include links starting from the strongest and stop this procedure as the total moisture transport volume exceeds the variable threshold ρ. The resulting edges then represent the backbone of the global moisture recycling network. In the main text, we have shown the results for a network where all edges together represent ρ = 25% of the total moisture transport. Here and in the SI figures, we add a sensitivity analysis for ρ = 20% and ρ = 30% and find that the results are stable for this broader range of total moisture volume thresholds.Network measures and motifsThe topology of an unweighted network is typically encoded in an adjacency matrix A with elements aij indicating if there exists an edge from node i to node j (aij = 1) or not (aij = 0). The degree k of a node i describes the number of adjacent edges pointing towards or away from node i. Hence, the in-degree is defined by25$${k}_{{{{{{mathrm{in}}}}}}}^{i}=mathop{sum }limits_{i=1}^{N}{a}_{ji}$$
    (5)
    and out-degree is defined by25$${k}_{{{{{{mathrm{out}}}}}}}^{i}=mathop{sum }limits_{i=1}^{N}{a}_{ij}.$$
    (6)
    To further analyze the topology of a network and, in particular, the local connectivity patterns, we study the presence of three motifs—the feed-forward loop, the neighboring loop, and the zero loop.The feed-forward loop (FFL) consists of three nodes, A, B, and C, where nodes A and C are directly connected via a detour over node B (intermediary node). Therefore, we have two different pathways that focus on node C. Hence, this motif can be referred to as a directed lens, due to its focused flow from two nodes on one singular and its purely directed linkage. This network motif has been studied in the context of tipping elements and has been proven to facilitate tipping cascades by lowering critical thresholds19. The zero loop (ZL) is made up of a bidirectional connection of two nodes. In contrast to the FFL, where node A does not receive feedback from node C, here, both nodes are dependent on each other without a preferred direction of network flow. This facilitates tipping to a much lesser degree than the FFL motif19. The neighboring loop (NBr) is an extension of the ZL. In this case, there is an additional node connected to one of the nodes of a zero loop. Hence, there is a two-step directionality in the motif, but in contrast to the FFL, this motif is characterized by reciprocity.We count the number of motifs a certain node is involved in the network. The number of FFLs is counted as the number that a certain node is a so-called “target” node. The target node is the node, on which the triangular structure of the motif is converging to, i.e., the node that has been referred to as node C above. The ZL is a symmetric motif for the two involved nodes. Therefore, the number of ZLs of a certain node in the network is counted directly as the number of bidirectional interactions of the inspected node. Lastly, the number of NBrs of a certain node is the number of being in the center of a neighboring loop. With this procedure, each node is characterized by its number of FFLs, ZLs, and NBrs (cf. ref. 19).Motif strength and their spatially aggregated differenceTo assess the presence of motifs and, in particular, their relative frequency, we first determine the numbers of FFLs, ZLs, and NBrs per node. Subsequently, we normalize these counts by the respective maximum to obtain the motif strength, which is shown for each network motif in Fig. S5. In Fig. S5a–c, we display the motifs for the global network, and in Fig. S5d–f for the land-to-land network.To specifically characterize the focus regions by means of the network topology, we evaluate which motifs dominate in which region. Consequently, we compute the difference of the motif strengths shown in Fig. S5 and reveal the patterns shown in Fig. 2. For spatially aggregated motif strength differences (Fig. 2c, d), we then compute the average of the respective values inside the highlighted boxes.Sensitivity to link threshold ρ
    The network analysis featured in the main text uses those moisture recycling edges that together represent ρ = 25% of all atmospheric moisture recycling on Earth. As we aimed to focus on the strongest moisture flows, we chose a threshold of ρ = 25% aggregating the strongest moisture transport pathways. This allows us to reveal the regions of strongest moisture connections, which are located in and close to the tropics, as we expected. Overall, the aim of this thresholding procedure is to utilize a network approach with unweighted edges but also take into account the large spread of moisture recycling strengths. To test the robustness of the results to the threshold value, we here show the same figures as above in the main text but with different thresholds ρ. Note that the error bars in Fig. 2 are based on the analysis featured in this part (the resulting differences using thresholds of ρ = 20% and ρ = 30%).Figures S6 and S7 show the in- and out-degree of the all-to-all and land-to-land network using a threshold of ρ = 20% (Fig. S6) and ρ = 30% (Fig. S7). Note that the color bar has been adjusted as the number of links differs substantially between the networks. The main difference between Figs. S6 and S7 is the greater emphasis on moisture recycling in the mid-latitudes in Fig. S7. This is a direct consequence of considering more, and thus also some weaker, links. Acknowledging this difference, we stress that especially the land-to-land patterns (Figs. S6c, d, S7c, d) are consistent. In particular, the four focus regions, as defined in the main text, stand out as the main global land-to-land moisture recycling hubs. To support this visual analysis of the in- and out-degree pattern, we furthermore compute the motif strengths for both network configurations for quantitative validation of the results.In line with the main text, we compare the FFL and ZL strength (see Fig. 2a–d). Not only the spatial patterns in our sensitivity analysis agree remarkably well with the results in the main text above, but also the focus regions remain basically the same (cf. Fig. S8 for ρ = 20% and Fig. S9 for ρ = 30% with Fig. 2). The only slight change is the shift towards a directed lens (spatially aggregated FFL and ZL strength difference) for the Amazon basin in the all-to-all network for increasing ρ (cf. Fig. S8c vs Fig. S9c vs Fig. 2c). We attribute the overproportional increase of the number of FFLs to those that include at least one oceanic grid cell to this noticeable shift. This underscores our characterization of the Amazon basin as a directed lens.The spatially aggregated FFL and NBr difference (Figs. S10, S11) is structurally the same as above, where we computed the FFL and ZL difference (see Figs. S8, S9). The spatial patterns and the aggregated values are robust against shifts of ρ. However, for the Amazon basin (AB), the number of FFLs increases overproportionally in the all-to-all network when we include more links in our analysis. In other words, the spatially aggregated FFL-strength for AB increases for higher thresholds ρ (cf. Figs. S10c, S11c and Fig. 2g).Sensitivity to the size of the focus regionsAnother aspect affecting the results is the spatial extent chosen as a focus region (i.e., the rectangles in Fig. 2). Varying the size of these rectangles affects the spatially aggregated measures. For all focus regions besides the Amazon Basin (AB), the values are not significantly affected by changing the rectangle size, as the values close to the focus regions are either coherently negative, as for the Congo Rainforest (CR) and the Indonesian Archipelago (IA), or close to zero (South Asia: SA). The AB is characterized by positive values (tendency to lensing), whereas the more southern parts along the Andes are marked by more negative (corridor/washing machine) values.Hence, we assess the stability of the results by using the spatial region covered by the Amazon rainforest (the extent of the Amazon rainforest is based on ref. 6) and compare them to the ones obtained by using the rectangle. The results featured in Fig. S12 indicate that only considering the rainforest-covered parts of the AB leads to similar or even more positive (lensing) values, confirming our conclusions that the Amazon rainforest region functions differently from the other focus regions.Notes on mapsThis paper makes use of perceptually uniform color maps developed by ref. 57. The underlying world maps have been created by cartopy58. More

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    Ecological transition and sustainable development: integrated statistical indicators to support public policies

    The link between SDGs and NRRPThe Italian National Recovery and Resilience Plan (NRRP) is part of the Next Generation EU (NGEU) program, the 750-billion-euro package, consisting of about half of grants, agreed by the European Union in response to the pandemic crisis. The main component of the NGEU program is the Recovery and Resilience Facility (RRF), which has a duration of six years, from 2021 to 2026, and a total size of €672.5 billion (€312.5 billion grants, the remaining €360 billion loans at subsidized rates).The Plan is developed around three strategic axes shared at European level: digitalization and innovation, ecological planning and social inclusion.The missions of the NRRP are as follows:

    Mission 1: Digitalization, innovation, competitiveness, culture and tourism

    Mission 2: Green revolution and ecological transition

    Mission 3: Infrastructure for sustainable mobility

    Mission 4: Education and research

    Mission 5: Cohesion and inclusion

    Mission 6: Health.

    With the aim of encouraging the debate on the use of sustainability indicators for monitoring the progress of the PNRR, a mapping of the correspondences between the 17 Sustainable Development Goals and the 6 Missions provided for by the NRRP is proposed (Fig. 1). In this way it is possible to identify the SDGs indicators that can be useful tools for achieving the missions of the NRRP.Figure 1Relationships between SDGs indicators and NRRP missions.Full size imageOf particular interest for the purposes of our work is Mission 2 (Green Revolution and Ecological Transition) of NRRP. It provides for investments and reforms for the circular economy and to improve waste management, strengthen separate collection infrastructure and modernize or develop new waste treatment plants. Substantial tax incentives are provided to increase the energy efficiency of buildings, to achieve progressive decarbonization, to increase the use of renewable energy sources. In addition, the Mission devotes resources to enhancing the capacity of electricity grids, their reliability, security, and flexibility (Smart Grid) and water infrastructure. The Mission also includes the issues of territorial security, with prevention and restoration interventions in the face of significant hydrogeological risks, the protection of green areas and biodiversity, and those related to the elimination of water and soil pollution, and the availability of water resources.The main components of this mission are:

    M2C1: Circular economy and sustainable agriculture

    M2C2: Renewable energy, hydrogen, grid, and sustainable mobility

    M2C3: Energy efficiency and upgrading of buildings

    M2C4: Protection of land and water resources.

    The analysis of Mission 2 (Green Revolution and Ecological Transition) finds ample space in the SDGs creating important interconnections between the different indicators present in the individual Goals and the objectives of the Mission itself.The SDGs indicators to support the NRRPThe SDGs indicators selected for the analysis of Mission 2 (Green Revolution and Ecological Transition) of the NRRP, are descripted in Table 1. We considered 13 indicators, selected from Goals 2, 6, 7, 11, 12 and 15 which may be of significant interest for the achievement of Mission 2. These indicators will then be attributed to the individual components of the mission.Table 1 Goal, indicators, measures e source of SDGs data.Full size tableThe indicators were chosen based on their relevance to the objectives of the mission and on the availability of data on a regional basis. For each main component we can use the following indicators:

    M2C1: Circular economy and sustainable agriculture:

    – Share of utilized agricultural area invested by organic crops

    – Growth rate of organic crops

    – Delivery of municipal waste to landfill.

    – Separate waste collection

    M2C2: Renewable energy, hydrogen, grid and sustainable mobility:

    M2C3: Energy efficiency and upgrading of buildings

    M2C4: Protection of land and water resources

    – Irregularities in water distribution

    – Sealing and soil consumption per capita

    – Soil sealing from artificial cover

    – Fragmentation of the natural and agricultural territory

    – Incidence of urban green areas on the urbanized surface of cities.

    The SDGs indicators at the level of territorial distribution in ItalyWe carry out a first analysis by territorial distribution for the different sets of main components of Mission 2.From a first analysis of the M2C1 indicators (Circular Economy and Sustainable Agriculture) it emerges that the share of agricultural area destined for organic crops is greater, especially in the Center and in the South of Italy. In 2019, the extent of organic farming in Italy reached 15.8% of the utilized agricultural area, almost double the EU average. However, the annual growth rate of the areas converted to organic farming or in the process of conversion (+ 1.8%) is the lowest since 2012 and is negative in the South, where for the second consecutive year there is a decrease (− 2.1% in the 2-year period 2017–2019). The dynamics of organic farming is an index of the spread of sustainable agricultural practices, which must be accompanied by measures that also consider the pressure on the environment generated by agriculture (Table 2).Table 2 M2C1 indicators—Circular economy and sustainable agriculture by territorial distribution (year 2019).Full size tableAlso, in the Central and Southern Italy area there is the greatest delivery of waste to landfills. Waste cycle management is crucial for living conditions and global health. The share of municipal waste landfilled is steadily decreasing at national level. In 2019, in fact, the part sent to landfill is equal to 20.9% of the total, down compared to the previous year (21.5%). The separate collection of municipal waste represents a further important step in view of the objective of reducing the amount of waste returned to the environment and, more specifically, of the delivery of waste to landfills. The 18.5 million tons of differentiated RU in 2019 represent 61.3% of national production, a share almost doubled compared to ten years ago and up from last year by 3.1 percentage points. Despite the evident progress, Italy is still marked by a considerable delay compared to the regulatory objectives, having not yet reached, in 2019, the target of 65% of separate collection planned for 2012. Critical issues are also observed in relation to the substantial territorial gaps, which disadvantage the Center and the South compared to the North, despite the distances have been reduced in recent years.
    Regarding the M2C2 Mission (Renewable Energy, Hydrogen, Network and Sustainable Mobility), national and international energy policies have been committed for years to the enhancement of renewable energy sources, with the aim of decarbonizing the economy and guaranteeing the commitments made in the field of climate change. In 2019, one year after the expiry of the objectives of the European Union’s Climate-Energy Package, fourteen Member States, including Italy, exceeded the target assigned at national level. In Italy, the overall share of energy from renewable sources in gross final consumption (CFL) of energy, equal to 18.2% (Table 3), a percentage slightly lower than the average of the EU27 (19.7%), is placed for the sixth consecutive year above the 17% target set for our country. However, for Italy to achieve the ambitious programs defined by the 2020 National Integrated Energy and Climate Plan, which set a 30% target for renewables by 2030, a further boost to production from renewable sources is necessary. The resources introduced by the National Recovery and Resilience Plan (NRRP) to achieve the “green revolution and ecological transaction” include significant investments in the energy field, focusing, among other components, on a further strengthening of the Sources from Renewable energy (FER).Table 3 M2C2 indicators—Renewable energy, hydrogen, network and sustainable mobility by territorial distribution (year 2019).Full size tableThe M2C3 Mission (Energy Efficiency and Upgrading of Buildings) devotes resources to enhancing the capacity of electricity grids, their reliability, safety, and flexibility (Smart Grid). Consistent with the objectives of reducing energy consumption pursued by European policies, the Italian figure for 2019 confirms the process of reducing Italian energy intensity, which marks a further contraction of 1.3%, reaching an overall negative balance compared to the last decade of 11.8%, with an average annual rate of change of − 1.2% (Table 4). The reduction in energy intensity is largely attributable to the effect of the measures in favor of energy efficiency, which, between 2011 and 2019, resulted in energy savings of 12 Mtoe/year, equal to 77% of the 2020 target set by the National Action Plan for Energy Efficiency 2017. A further acceleration of energy efficiency is expected, in the coming years, because of the investment plan envisaged by the NRRP, also linked to the redevelopment of the public and private building stock. At the sectoral level, the reduction in energy intensity is driven by improvements in industry, which, despite the slight increase in the last year, in 2019, with 92 toes per million euros, shows a decrease compared to 2009 of 17%, with an average annual rate of change of − 1.8%.Table 4 M2C3 indicators—Energy efficiency and requalification of buildings by territorial distribution (year 2019).Full size tableThe M2C4 Mission (Protection of the territory and water resources) also includes the issues of territorial safety, with prevention and recovery interventions, the protection of green areas and those related to the elimination of water and soil pollution.Italy is among the European countries of the Mediterranean area that use groundwater, springs and wells the most; these represent the most important resource of fresh water for drinking water use on the Italian territory (84.8% of the total withdrawn). The efficiency of municipal drinking water distribution networks has been steadily deteriorating since 2008 for more than half of the regions. The share of families who complain of irregularities in the water supply service in their home is stable (equal to 8.6% in 2019) with more accentuated values in the Center and South of Italy (Table 5).Table 5 M2C4 indicators—Protection of land and water resources by territorial distribution (year 2019).Full size tableLand degradation, understood as loss of ecological functionality, is monitored through the dynamics of land consumption, which Italy has committed to zero by 2030 with the National Strategy for Sustainable Development (2017). The “consumed” soil is that occupied by urbanization and made impermeable by artificial roofing (soil sealing). Excessive fragmentation of open spaces, however, is also a factor of degradation, since the barriers made up of buildings and infrastructures interrupt the continuity of ecosystems, making even unoccupied but not large enough spaces ecologically inert and unproductive. Moreover, in a fragile territory such as Italy, land consumption is also a significant factor of hydrogeological risk and deterioration of the landscape. The index of sealing and land consumption per capita in 2019 increases for the fifth consecutive year, resulting in 357 m2 per inhabitant. The soil sealed by artificial covers is equal to 7.1% of the national territory (8.5% in the North, 6.7% in the Center, 5.9% in the South).According to Ispra estimates, 44.3% of Italy’s natural and agricultural land has a high or very high degree of fragmentation. A joint representation of the variations in fragmentation and soil sealing over the last two years summarizes recent trends in land consumption and their impact on the environment and landscape.A further objective for 2030 is to provide universal access to safe, inclusive, and accessible public green spaces, for women and children, the elderly, and people with disabilities. In 2019 the incidence of urban green areas on the urbanized surface of cities is equal to 8.5% in Italy with slightly higher values in the North and less elevated in the South. More

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    Investigation of the spermathecal morphology, reproductive strategy and fate of stored spermatozoa in three important thysanopteran species

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    Quantifying the impacts of land cover change on gross primary productivity globally

    GPP dataAs our primary productivity product we used the GOSIF GPP dataset21 which utilizes the linear relationship between GPP and remotely-sensed SIF34. GOSIF GPP is available globally at 0.05° spatial resolution for the period 2000–2021, with the period 2001–2015 selected here (for a short summary of all datasets used in this study see Supplementary Table 3). GOSIF GPP is based on a gridded SIF product (GOSIF)34 which uses MODIS enhanced vegetation index and meteorological data for spatial scaling and is trained with millions of SIF observations from the coarser-resolution Orbiting Carbon Observatory-235. The global coverage of GOSIF and the close relationship between SIF and GPP allow for an independent assessment of how land cover changes affect GPP in different regions around the world. For instance, SIF has been shown to capture the high GPP in the US Corn Belt derived from flux towers, while ecosystem models underestimated it36. While GPP can thus be empirically estimated from satellite SIF observations relatively reliably (even though some assumptions like the linear GPP–SIF relationship and its universality across biomes are still debated20,37,38,39), the calculation of NPP needs additional assumptions of autotrophic respiration. Therefore, we focused our study on GPP, but we included an NPP product in our uncertainty analysis. In addition to that, to account for the challenges and uncertainties in global GPP estimates we included four alternative GPP products in our sensitivity analysis (see below).Land cover mappingGridded land cover was derived from ESA-CCI22, a global land cover product designed for climate science. ESA-CCI is available at 300 m spatial resolution for the 1992–2020 period (https://cds.climate.copernicus.eu/). We first classified ESA-CCI land covers to forests, grasslands, and croplands according to IPCC classification: classes 50–100, 160, 170 forests (2,022,283 grid cells); classes 110 and 130 grasslands (509,297 grid cells); classes 10–40 croplands (950,025 grid cells). We focus on these three major land cover types to facilitate our analysis. We then converted the resulting map to 0.05° resolution by determining the prevalent (i.e., mode) land cover for each grid cell using the aggregate function from the raster package40 and only included grid cells in our training data in which the prevalent land cover was constant over the period 2001–2015. Other classes (e.g., cropland/natural vegetation mosaics) and grid cells where the land cover changed over the 2001–2015 period were not used for the RF training.Random forestsRF is a popular and efficient supervised machine learning technique which can be applied for classification and regression problems41. While complex, it is still easier to interpret compared to other machine learning methods such as Artificial Neural Networks. It has recently been applied to a wide range of ecological research questions, including the prediction of food42 and bioenergy43 crop yields, potential natural vegetation31, forest aboveground biomass44, soil respiration45, and soil carbon emissions from land-use change5 and is thus well suited for our approach. The “Forests” refer to a number of individual decision trees. For each tree, a random sample of the training data is selected and split multiple times based on a random subset of variables from which the one minimizing the weighted variance is selected by the algorithm. Model performance is computed directly on out-of-bag (OOB) data which is randomly omitted from the training data (36.8% of all grid cells). When RF is applied to new data, a weighted prediction of each individual decision tree contributes to the overall prediction. Variance in the individual trees, e.g., by selecting random subsets of the observations and random variables at each node improves the overall RF predictive skill. Model training and prediction were done using the R ranger package46. After initial testing (see Supplementary Fig. S11) we decided to set the number of individual decision trees to 800 and the number of variables to possibly split at in each node to 10. As the good evaluation measures of RF algorithms can be related to spatial autocorrelation24 we also tested a coordinate-only model and performed a leave-one-out cross validation including spatial buffers (Supplementary Discussion 2, Supplementary Fig. S3). Due to the large computational effort we reduced the number of decision trees to 100 for the buffered leave-one-out cross validation.Predictor variablesWe predicted forest, grassland, and cropland potential GPP using the following 20 predictor variables in our RF algorithm: mean annual surface temperature (Tmean), mean diurnal temperature range (Tdiurnal), temperature seasonality (Tseason; standard deviation), minimum temperature of the coldest month (Tmin), annual temperature range (Tannual), mean temperature of the warmest quarter (Twarmest), mean annual precipitation (Pmean), precipitation seasonality (Pseason; coefficient of variation), precipitation of the wettest quarter (Pwettest), precipitation of the driest quarter (Pdriest), precipitation of the warmest quarter (Pwarmest), mean annual solar radiation (SR), growing degree days (GDD), relative humidity (RH), soil clay content (Clay), elevation (EL), nitrogen deposition (Ndep), nitrogen fertilization (NF), pesticide application (Pest), and gross domestic product (GDP; a proxy for agricultural management input other than NF and Pest). Overall Tmean, Tannual, and Pmean were the most important predictor variables (see Supplementary Discussion 3 and Fig. S12). We also tested other predictors (including additional bioclimatic variables, soil pH, irrigation, or phosphate fertilization) but found only negligible improvements in RF evaluation metrics and hence decided to restrict our analysis to the 20 predictors mentioned above.Climate variables were taken from the CHELSA dataset47,48, remapped to 0.05° spatial resolution using the aggregate function from the raster package40. To only include years overlapping with our GPP data we used the CHELSA time-series data for the 2001–2013 period if available and 1979–2013 climatologies elsewise. Clay was derived from the Regridded Harmonized World Soil Database v1.249. Ndep was taken from ISIMIP2b50, bilinear remapped from 0.5° to 0.05° spatial resolution using Climate Data Operators33. Elevation was obtained from WorldClim51. NF and Pest were derived from country-specific FAO data (e.g., https://ourworldindata.org/grapher/pesticide-use-per-hectare-of-cropland), i.e., we used the same value for all grid cells in a country. GDP was obtained from ref.52.Suitable areaFor the comparison of potential forest, grassland, and cropland GPP in Fig. 1g–i we only included grid cells suitable for all three land cover types. For forests, we assumed forest cover possible if the grid cell is currently forested (e.g., all grid cells of our forest training data) or if the potential natural forest cover exceeds 36.3%. This threshold represents the 5th percentile of all currently forested grid cells. Potential natural forest cover was derived from a potential natural vegetation map, available for 20 biomes at 0.00833° spatial resolution31. To convert these biomes into potential natural forest cover we assumed 100% forest cover for the ten forest biomes and 30% forest cover for tropical savannah. Other biomes were not considered. We then aggregated the map to 0.05° spatial resolution by computing the mean of 36 grid cells using the aggregate function form the raster package40 (see Supplementary Fig. S5 for the resulting map). For grasslands and croplands, we computed the 5th percentile of Tmean and Pmean in the training data (− 9.9 °C and 165 mm for grasslands and 2.7 °C and 295 mm for croplands, respectively) and removed all grid cells below those thresholds, assuming these areas to be too cold or too dry for the respective land cover type. Finally, we calculated the land cover with the highest potential GPP for all overlapping grid cells.Sensitivity analysisTo explore the sensitivity and uncertainty of our RF approach we repeated our prediction using different input datasets, potential forest cover, and machine-learning approaches. The importance of the underlying potential forest map was estimated by replacing our potential forest map (Supplementary Fig. S5) by the LUH2 potential forest map (Supplementary Fig. S13)23. To explore the dependency on the land cover product we repeated our RF prediction using the spatially aggregated MODIS land cover map (MCD12C1; IGBP scheme), available at 0.05° spatial resolution53. We classified grid cells of classes 1, 2, 3, 4, 5, (all forests), 8 (woody savannahs) and 9 (savannahs) as forest. Classes 8 and 9 were included in forest because otherwise forest cover would be underestimated in the temperate and boreal zone. Class 10 was classified as grassland and class 12 as cropland. A comparison of ESA-CCI with MODIS reveals a substantially larger cropland area in ECA-CCI but a smaller grassland area (Supplementary Fig. S14).The sensitivity to the climate product was tested by repeating our analysis using predictor variables from the WorldClim climatologies (1970–2000)51, aggregated from 30 s to 0.05° spatial resolution using the aggregate function from the raster package40. In contrast to CHELSA, growing degree days and relative humidity were not available from WorldClim but we included water vapour pressure as additional predictor.We also tested four alternative global GPP products. The vegetation photosynthesis model (VPM) product, available for the period of interest at 0.05° spatial resolution, is based on improved light use efficiency theory and is driven by remotely sensed datasets and reanalysis climate data and land cover classification which also distinguishes C3 vs. C4 photosynthesis pathways54. The second product is derived from remote sensing considering radiation and canopy conductance limitations on GPP and is available at 0.05° resolution for the 2001–2012 period55. Land cover is not an input variable. The third product, FLUXCOM, uses machine learning to scale FLUXNET site GPP to the globe56,57. FLUXCOM is available at 0.0833° resolution and was conservative remapped to 0.05° using Climate Data Operators33 meaning that the GPP of different land cover types might be mixed in regions with heterogeneous land cover patterns. The forth product is the MODIS MOD17A3 GPP product58, available for the 2001–2013 period and aggregated to 0.05° resolution using the raster package40. It is derived from meteorological data, fraction of absorbed photosynthetic active radiation/leaf area index, and land cover. As there is also a MOD17A3 NPP product available we additionally conducted a prediction for potential NPP. The MOD17A3 NPP product is calculated as GPP minus maintenance and growth respiration estimated from allometric relationships linking daily biomass and annual growth of plant tissues to leaf area index58. This leads to additional uncertainty compared to the MOD17A3 GPP product.To test the effect of an alternative RF algorithm we repeated our prediction with the RF algorithm from the Python scikit-learn library59 using the same number of decision trees (800). Additionally, we tested another machine-learning technique, a deep neural network (DNN), using the PyTorch library60. We selected 10 linear layers with 5 times alternating 128 and 256 nodes and a sigmoid output function. All layers were connected using the rectified linear unit activation function. We used the adamW optimizer with 0.0003 learning rate and 2000 epochs of training. To prevent overfitting, we included a 10% dropout after the 7th layer. Lastly, we included a very simple estimate of the most productive land cover based on the nearest neighbour using scikit-learn’s BallTree implementation together with the Haversine formula. For each grid cell we searched for the nearest forest, grassland, and cropland grid cell and assigned the respective GPP also to this grid cell. We thus assumed that environmental conditions are more or less identical in these grid cells, which might be a reasonable assumption for many locations but less reliable in complex terrain or in large homogeneous regions like the central Amazon rainforest where the nearest cropland/grassland grid cell might be located far away.Land-use change scenariosTo estimate the effects of historical and potential future land cover changes on global GPP we applied LUH2 scenarios23 which also serve as input data for ESMs participating in CMIP6. Land-use changes over the historical period are based on the HYDE reconstruction3, while future projections were developed by different Integrated Assessment Models combining various assumptions of socio-economic behaviour (SSPs) with climate mitigation targets (RCPs). Annual fractions for the two land cover classes cropland (sum of 5 crop types) and managed grassland (sum of pasture and rangeland) were available for each scenario at 0.25° resolution (https://luh.umd.edu/). We converted to 0.05° resolution assuming the same land cover fractions for all 25 grid cells around the LUH2 grid cells. We considered the following land cover transitions: forest to managed grassland, forest to cropland, and natural grassland to cropland (and reverse transitions for future scenarios). Transitions in areas suitable for only two land cover types were also included. Conversions of natural grasslands to managed grasslands were assumed not to affect productivity. We assumed the original land cover of a grid cell to be either forest (i.e., potential forest cover  > 36.3%) or natural grassland and accordingly multiplied the converted areas by the differences in potential GPP derived from our RF approach. Our broad forest definition including open tree cover (see above) and the fact that we assumed a change from 100 to 0% forest area in deforested grid cells results in a total historical deforestation area substantially larger than estimated in a recent study (2.4 Mkm2 vs. 1.6 Mkm2)61. These assumptions, however, do not impair our GPP estimate as our approach implicitly accounts for gradients in forest productivity (open forests tend to have lower GPP than closed forests). To test the sensitivity of the resulting GPP reduction we also applied the potential GPP maps from our uncertainty analysis to historical land-use changes (Supplementary Fig. S6). For future land cover changes we investigated changes over the 2015–2100 period for all available LUH2 scenarios: SSP1-1.9, SSP2-2.6, SSP4-3.4, SSP5-3.4, SSP2-4.5, SSP4-6.0, SSP3-7.0, and SSP5-8.5. Land-use activities in these scenarios range from large-scale deforestation (e.g., SSP3-7.0) to reforestation (e.g., SSP1-1.9) (Supplementary Fig. S7).Earth System ModelsWe compared the potential GPP estimated by our RF algorithm to simulations of eight ESMs participating in CMIP6 (CESM2-CLM562, CNRM-ESM2.1-Surfex 8.0c63, EC-Earth3-Veg-LPJ-GUESSv464, GFDL-ESM4-GFDL-LM4.165, IPSL-CM6A-LR-ORCHIDEEv2.066, MIROC-ES2L-MATSIRO6.0 + VISIT-e ver.1.067, MPI-ESM1-2-LR-JSBACH3.2068, UKESM1-0-LL-JULES-ES-1.069) with an explicit representation of natural vegetation and at least one agricultural land cover class (cropland or managed grassland) in their vegetation sub-model. We selected these ESMs so that all vegetation models implemented in more than one ESM were represented only once (e.g., the JSBACH vegetation model is a component of both MPI-ESM1-2-LR and AWI-ESM). For each ESM, the variable gppLut was downloaded from the CMIP6 archive (https://esgf-data.dkrz.de/search/cmip6-dkrz/) for the historical simulations. These files contain simulated GPP for natural vegetation, pasture, and cropland for which we calculated the 2001–2014 mean (2014 is the last year of the historical period). ESMs use fractional land covers for each grid cell, meaning that climatic drivers are inherently the same for all land cover types within a grid cell and simulated productivities can therefore be directly compared. As ESMs differ in their spatial resolution we bilinear remapped all output to 0.05° resolution using Climate Data Operators33 to allow for a fair comparison across models. To assess the sensitivity of our results to the interpolation method we also tested conservative remapping which results in slightly different maps (Supplementary Fig. S15) and usually larger model biases (Supplementary Table 2). In addition, ESMs differ in where they simulate forests in natural vegetation areas, and therefore we removed all grid cells from the comparison where at least one ESM simulated no tree productivity/cover/biomass in order to avoid comparing the GPP of natural grasslands to managed grasslands. We provide maps based on the original output for each ESM in Supplementary Fig. S10.FLUXNET dataWe compared our predictions of potential GPP to FLUXNET Tier 1 eddy covariance measurements (Supplementary Fig. S16)70. We included all forest, woody savannah (classified as forest), grassland and cropland sites21 which were located in suitable areas for the respective land cover. Mean GPP was calculated as the mean of the GPP estimates based on the night-time (GPP_NT_VUT_REF) and day-time (GPP_DT_VUT_REF) partitioning method. As some sites only had a few years of data, all available years were considered (i.e., site mean GPP was calculated for a different time period than 2001–2015). Comparisons were made with the potential GPP in the respective grid cell in which the site was located (i.e., not calibrated to site conditions). More

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    Impact of host age on viral and bacterial communities in a waterbird population

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