Population density per pixel at 100 metre resolution. WorldPop provides estimates of numbers of people residing in each 100x100m grid cell for every low and middle income country. Through ingegrating cencus, survey, satellite and GIS datasets in a flexible machine-learning framework, high resolution maps of population counts and densities for 2000-2020 are produced, along with accompanying metadata.

DATASET: Alpha version 2010 and 2015 estimates of numbers of people per grid square, with national totals adjusted to match UN population division estimates (http://esa.un.org/wpp/) and remaining unadjusted.

REGION: Africa

SPATIAL RESOLUTION: 0.000833333 decimal degrees (approx 100m at the equator)

PROJECTION: Geographic, WGS84

UNITS: Estimated persons per grid square

MAPPING APPROACH: Land cover based, as described in: Linard, C., Gilbert, M., Snow, R.W., Noor, A.M. and Tatem, A.J., 2012, Population distribution, settlement patterns and accessibility across Africa in 2010, PLoS ONE, 7(2): e31743.

FORMAT: Geotiff (zipped using 7-zip (open access tool): www.7-zip.org)

FILENAMES: Example - AGO10adjv4.tif = Angola (AGO) population count map for 2010 (10) adjusted to match UN national estimates (adj), version 4 (v4). Population maps are updated to new versions when improved census or other input data become available.

India data available from WorldPop here.

Census data reveals that population density varies noticeably from area to area. Small area census data do a better job depicting where the crowded neighborhoods are. In this map, the yellow areas of highest density range from 30,000 to 150,000 persons per square kilometer. In those areas, if the people were spread out evenly across the area, there would be just 4 to 9 meters between them. Very high density areas exceed 7,000 persons per square kilometer. High density areas exceed 5,200 persons per square kilometer. The last categories break at 3,330 persons per square kilometer, and 1,500 persons per square kilometer.This dataset is comprised of multiple sources. All of the demographic data are from Michael Bauer Research with the exception of the following countries:Australia: Esri Australia and MapData ServicesCanada: Esri Canada and EnvironicsFrance: Esri FranceGermany: Esri Germany and NexigaIndia: Esri India and IndicusJapan: Esri JapanSouth Korea: Esri Korea and OPENmateSpain: Esri España and AISUnited States: Esri Demographics

Facebook and Columbia University - CIESIN provide the High Resolution Settlement Layer as the world's most accurate population datasets. More info can be found here: https://dataforgood.fb.com/tools/population-density-maps/

These maps are the distribution of human population spanning Pakistan and India. Each of the 13 TIFF files is a 10 x 10 degree tile (the lower latitude coordinate and longitude coordinates are in the file name). A VRT file is also included.

Soil is a key natural resource that provides the foundation of basic ecosystem services. Soil determines the types of farms and forests that can grow on a landscape. Soil filters water. Soil helps regulate the Earth's climate by storing large amounts of carbon. Activities that degrade soils reduce the value of the ecosystem services that soil provides. For example, since 1850 35% of human caused green house gas emissions are linked to land use change. The Soil Science Society of America is a good source of of additional information.Bulk density is an important property of soil. Soil is a mixture of mineral particles, organic material and open spaces known as pores. The size and distribution of the pores affect how water is stored and how nutrients move through the soil. When a soil is compacted it looses pore space and bulk density increases resulting in lower water storage and higher runoff.Dataset SummaryThis layer provides access to a 30 arc-second (roughly 1 km) cell-sized raster with attributes related to the density of soil derived from the Harmonized World Soil Database v 1.2. The values in this layer are for the dominant soil in each mapping unit (sequence field = 1).Attribute values for topsoil (0-30 cm) and subsoil (0-100 cm) are provided for bulk density (derived from available analyzed data) and reference bulk density (statistical estimate based on soil texture). The data are in units of kg/dm3. Topsoil Reference Bulk DensityTopsoil Bulk Density Subsoil Reference Bulk DensitySubsoil Bulk DensityThe layer is symbolized with the Topsoil Bulk Density field.The document Harmonized World Soil Database Version 1.2 provides more detail on the difference between bulk density and reference bulk density.Other attributes contained in this layer include:Soil Mapping Unit Name - the name of the spatially dominant major soil groupSoil Mapping Unit Symbol - a two letter code for labeling the spatially dominant major soil group in thematic mapsData Source - the HWSD is an aggregation of datasets. The data sources are the European Soil Database (ESDB), the 1:1 million soil map of China (CHINA), the Soil and Terrain Database Program (SOTWIS), and the Digital Soil Map of the World (DSMW).Percentage of Mapping Unit covered by dominant componentObstacles to Roots - Depth to obstacles to roots in 6 classes. Only in the European Soil Database, not available for other regions.More information on the Harmonized World Soil Database is available here.Other layers created from the Harmonized World Soil Database are available on ArcGIS Online:World Soils Harmonized World Soil Database – ChemistryWorld Soils Harmonized World Soil Database - Exchange CapacityWorld Soils Harmonized World Soil Database – GeneralWorld Soils Harmonized World Soil Database – HydricWorld Soils Harmonized World Soil Database – TextureThe authors of this data set request that projects using these data include the following citation:FAO/IIASA/ISRIC/ISSCAS/JRC, 2012. Harmonized World Soil Database (version 1.2). FAO, Rome, Italy and IIASA, Laxenburg, Austria.What can you do with this layer?This layer is suitable for both visualization and analysis. It can be used in ArcGIS Online in web maps and applications and can be used in ArcGIS Desktop.This layer has query, identify, and export image services available. This layer is restricted to a maximum area of 16,000 x 16,000 pixels - an area 4,000 kilometers on a side or an area approximately the size of Europe. The source data for this layer are available here.This layer is part of a larger collection of landscape layers that you can use to perform a wide variety of mapping and analysis tasks.The Living Atlas of the World provides an easy way to explore the landscape layers and many other beautiful and authoritative maps on hundreds of topics.Geonet is a good resource for learning more about landscape layers and the Living Atlas of the World. To get started follow these links:Living Atlas Discussion GroupSoil Data Discussion GroupThe Esri Insider Blog provides an introduction to the Ecophysiographic Mapping project.

The territories of Pakistan and India are mostly covered by the non-political blocks AS42 through AS50, going roughly from West to East. Please see the attached map of these non-political boundary blocks.

In 2022, the union territory of Delhi had the highest urban population density of over ** thousand persons per square kilometer. While the rural population density was highest in union territory of Puducherry, followed by the state of Bihar.

Analysis of ‘Indian Census Data with Geospatial indexing’ provided by Analyst-2 (analyst-2.ai), based on source dataset retrieved from https://www.kaggle.com/sirpunch/indian-census-data-with-geospatial-indexing on 13 February 2022.

--- Dataset description provided by original source is as follows ---

Dataset Description:

• This dataset has population data of each Indian district from 2001 and 2011 censuses.
• The special thing about this data is that it has centroids for each district and state.
• Centroids for a district are calculated by mapping border of each district as a polygon of latitude/longitude points in a 2D plane and then calculating their mean center.
• Centroids for a state are calculated by calculating the weighted mean center of all districts that constitutes a state. The population count is the weight assigned to each district.

Example Analysis:

• The complete code for calculating the centroids and web scraping for the data is shared on GitHub.

• The purpose of this project was to map population density center for each state.

• You can also read about the complete project here: https://medium.com/@sumit.arora/plotting-weighted-mean-population-centroids-on-a-country-map-22da408c1397

Output Screenshots: Indian districts mapped as polygons https://i.imgur.com/UK1DCGW.png" alt="Indian districts mapped as polygons">

Mapping centroids for each district https://i.imgur.com/KCAh7Jj.png" alt="Mapping centroids for each district">

Mean centers of population by state, 2001 vs. 2011 https://i.imgur.com/TLHPHjB.png" alt="Mean centers of population by state, 2001 vs. 2011">

National center of population https://i.imgur.com/yYxE4Hc.png" alt="National center of population">

--- Original source retains full ownership of the source dataset ---

Population density in the Ghataprabha (K3) sub-basin area. The dataset, calculated as number of people per hectare, is derived from the WorldPop datasets (https://www.worldpop.org/)

Population density in the Malaprabha (K4) sub-basin area. The dataset, calculated as number of people per hectare, is derived from the WorldPop datasets (https://www.worldpop.org/)

Monaco led the ranking for countries with the highest population density in 2024, with nearly 26,000 residents per square kilometer. The Special Administrative Region of Macao came in second, followed by Singapore. The world’s second smallest country Monaco is the world’s second-smallest country, with an area of about two square kilometers and a population of only around 40,000. It is a constitutional monarchy located by the Mediterranean Sea, and while Monaco is not part of the European Union, it does participate in some EU policies. The country is perhaps most famous for the Monte Carlo casino and for hosting the Monaco Grand Prix, the world's most prestigious Formula One race. The global population Globally, the population density per square kilometer is about 60 inhabitants, and Asia is the most densely populated region in the world. The global population is increasing rapidly, so population density is only expected to increase. In 1950, for example, the global population stood at about 2.54 billion people, and it reached over eight billion during 2023.

Population density in the Middle Krishna (K2) sub-basin area. The dataset, calculated as number of people per hectare, is derived from the WorldPop datasets (https://www.worldpop.org/)

Additional file 1: Basic Information of India. Table S1. List of Indian States and Union Territories. Figure S1. Map of Indian States and Union Territories. Figure S2. Map of Indian population density. Figure S3. Averaged annual rainfall map of India (2013-2016). The red arrows are monsoon move directions during summer.

Maps with wind speed, wind rose and wind power density potential in India. The GIS data stems from the Global Wind Atlas (http://globalwindatlas.info/). GIS data is available as JSON and CSV. The second link provides poster size (.pdf) and midsize maps (.png).

Snapshot img

Digital Map Market Size 2025-2029

The digital map market size is forecast to increase by USD 31.95 billion at a CAGR of 31.3% between 2024 and 2029.

The market is driven by the increasing adoption of intelligent Personal Digital Assistants (PDAs) and the availability of location-based services. PDAs, such as smartphones and smartwatches, are becoming increasingly integrated with digital map technologies, enabling users to navigate and access real-time information on-the-go. The integration of Internet of Things (IoT) enables remote monitoring of cars and theft recovery. Location-based services, including mapping and navigation apps, are a crucial component of this trend, offering users personalized and convenient solutions for travel and exploration. However, the market also faces significant challenges.
Ensuring the protection of sensitive user information is essential for companies operating in this market, as trust and data security are key factors in driving user adoption and retention. Additionally, the competition in the market is intense, with numerous players vying for market share. Companies must differentiate themselves through innovative features, user experience, and strong branding to stand out in this competitive landscape. Security and privacy concerns continue to be a major obstacle, as the collection and use of location data raises valid concerns among consumers.

What will be the Size of the Digital Map Market during the forecast period?

Explore in-depth regional segment analysis with market size data - historical 2019-2023 and forecasts 2025-2029 - in the full report.
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In the market, cartographic generalization and thematic mapping techniques are utilized to convey complex spatial information, transforming raw data into insightful visualizations. Choropleth maps and dot density maps illustrate distribution patterns of environmental data, economic data, and demographic data, while spatial interpolation and predictive modeling enable the estimation of hydrographic data and terrain data in areas with limited information. Urban planning and land use planning benefit from these tools, facilitating network modeling and location intelligence for public safety and emergency management.

Spatial regression and spatial autocorrelation analyses provide valuable insights into urban development trends and patterns. Network analysis and shortest path algorithms optimize transportation planning and logistics management, enhancing marketing analytics and sales territory optimization. Decision support systems and fleet management incorporate 3D building models and real-time data from street view imagery, enabling effective resource management and disaster response. The market in the US is experiencing robust growth, driven by the integration of Geographic Information Systems (GIS), Global Positioning Systems (GPS), and advanced computer technology into various industries.

How is this Digital Map Industry segmented?

The digital map industry research report provides comprehensive data (region-wise segment analysis), with forecasts and estimates in 'USD million' for the period 2025-2029, as well as historical data from 2019-2023 for the following segments.

Application

  Navigation
  Geocoders
  Others


Type

  Outdoor
  Indoor


Solution

  Software
  Services


Deployment

  On-premises
  Cloud


Geography

  North America

    US
    Canada


  Europe

    France
    Germany
    UK


  APAC

    China
    India
    Indonesia
    Japan
    South Korea


  Rest of World (ROW)

By Application Insights

The navigation segment is estimated to witness significant growth during the forecast period. Digital maps play a pivotal role in various industries, particularly in automotive applications for driver assistance systems. These maps encompass raster data, aerial photography, government data, and commercial data, among others. Open-source data and proprietary data are integrated to ensure map accuracy and up-to-date information. Map production involves the use of GPS technology, map projections, and GIS software, while map maintenance and quality control ensure map accuracy. Location-based services (LBS) and route optimization are integral parts of digital maps, enabling real-time navigation and traffic data.

Data validation and map tiles ensure data security. Cloud computing facilitates map distribution and map customization, allowing users to access maps on various devices, including mobile mapping and indoor mapping. Map design, map printing, and reverse geocoding further enhance the user experience. Spatial analysis and data modeling are essential for data warehousing and real-time navigation. The automotive industry's increasing adoption of connected cars and long-term evolution (LTE) technologies have fueled the demand for digital maps. These maps enable driver assistance app

The R-factor serves as a climatological index, reflecting the intensity and kinetic energy of rainfall and playing a pivotal role in evaluating the impacts of sheet and rill erosion.

The InRESatE dataset introduces rainfall erosivity data, estimated from high-resolution IMERG Final Run 30-minute precipitation records spanning 2001 to 2018. Additionally, the dataset includes erosivity density values and calibrated parameters for a daily rainfall erosivity model tailored to the study area.

Our analysis reveals an annual rainfall erosivity range of 77 to 20,662 MJ mm ha−1h−1 yr−1, with the summer monsoons accounting for approximately 85% of this yearly erosivity. While our erosivity map closely aligns with previous versions of rainfall erosivity datasets based on sparse gauge data, it registers marginally lower values in regions experiencing extreme rainfall. Erosivity density maps highlight erosion and landslide-prone zones, particularly in India's Northeast and Western Ghats regions, attributed to intense summer monsoon rainfall.

Article Citation:

Das, Subhankar, Manoj Kumar Jain, and Vivek Gupta. "A step towards mapping rainfall erosivity for India using high-resolution GPM satellite rainfall products." Catena 212 (2022): 106067. https://doi.org/10.1016/j.catena.2022.106067

ABSTRACTNatural hazards like landslides, earthquakes, and floods are a major deterrent to the development of mountain regions of the world. Recently, there has been a rise in the number of landslides in Western Ghats, India. Several factors cause landslides and they depend on the local geo-environmental set-up of the region. This study attempts to map the spatial distribution of landslide hazard and analyze the risk for a typical hill town, Kodaikkanal in the Western Ghats, India, facing rapid urbanization and infra-structure growth, using a weighted linear combination model in a Geographic Information System platform. Landslides in the region are triggered by rainfall and it is fairly uniform throughout the area. Hence, the susceptibility map is used as the hazard map. Validation of the weighted linear combination model using landslide hazard index shows that landslide density increases with the hazard class. The risk assessment matrix (RAM) is used to evaluate risk based on the land use and landslide hazard category. The land use map is reclassified by assigning damage potential for each land use feature. The risk map is classified into low, moderate, and high risk categories. Suitable control measures are suggested for various risk categories.

This data package includes spatial environmental and social layers for Shivamogga District, Karnataka, India that were considered as potential predictors of patterns in human cases of Kyasanur Forest Disease (KFD). KFD is a fatal tick-borne viral haemorrhagic disease of humans, that is spreading across degraded forest ecosystems in India. The layers encompass a range of fifteen metrics of topography, land use and land use change, livestock and human population density and public health resources for Shivamogga District across 1km and 2km study grids. These spatial proxies for risk factors for KFD that had been jointly identified between cross-sectoral stakeholders and researchers through a co-production approach. Shivamogga District is the District longest affected by KFD in south India. The layers are distributed as 1km and 2km GeoTiffs in Albers equal area conic projection. For KFD, spatial models incorporating these layers identified characteristics of forest-plantation landscapes at higher risk for human KFD. These layers will be useful for modelling spatial patterns in other environmentally sensitive infectious diseases and biodiversity within the district.

Methods Processing of environmental predictors of Kyasanur Forest Disease distribution

This file details the sources and processing of environmental predictors offered to the statistical analysis in the paper. All processing was performed in the raster package [1] of the R program [2] unless otherwise specified, with function names as specified below.

Topography predictors

Elevation data was extracted in tiles from Shuttle Radar Topography Mission data version 4 [3] an original resolution of 0.000833 degrees Latitude and Longitude resolution (approximately 90m by 90m grid cells). Tiles were mosaicked across the study region using the merge function. A slope value for each pixel was calculated (in degrees) using the terrain function of the raster package, and a focal window of 3 by 3 cells. Both the resulting elevation and slope rasters were cropped to the administrative boundaries of the Shivamogga District (raster package: crop function) and re-projected to an equal area projection (Albers equal area conic projection) using the projectRaster function (method=”bilinear”). Mean elevation and slope values were then calculated across the study 1km and 2km grid cells, using the aggregate function to average values across the appropriate number of ~90m grid cells and then the resample function to align the resulting grid to the study grids.

Landscape predictors

Metrics of the current availability (and fragmentation) of forest, agricultural and built-up land use types as well as that of water-bodies were extracted from the MonkeyFeverRisk Land Use Land Cover map of Shimoga. The latter was produced from classification of earth observation data from 2016 to 2018 using the methods described in the Supplementary information S3 file of the paper linked to this dataset. The LULC map had an original grid square resolution of 0.000269 degrees Latitude and Longitude resolution (or 30m x 28m grid cells) and nine different LULC classes. It was cropped to the administrative boundaries of the Shimoga District (raster package: crop function) and re-projected to the equal area projection (Albers equal area conic projection) using the projectRaster function (method=”ngb” for categorical data). The agriculture and fallow land classes were combined before landscape analysis (due to the difficulty of separating them accurately in the classification process).

An algorithm was developed in R to identify which of the pixels in the LULC map coincided with each 1km and 2km grid cell of the study area. The ClassStat function of the SDM Tools package [4] was used to calculate the proportional area of each 1km or 2km grid cell landscape that was made up of a particular land class, as well patch density and edge density metrics for the forest classes as indicators of fragmentation and forest-agriculture interface habitat respectively (Fig. S2B). The proportional area values (pi) of the n different forest classes (wet evergreen forest, moist deciduous forest, dry deciduous forest and plantation) were used to calculate an index of forest type diversity per grid cell as follows, after Shannon & Weaver (1949) [5]:

H'= -1npi(lognpi)

Metrics of longer term forest changes in Shimoga since 2000 were derived from a global product by Hansen et al. (2013) [6] available at a spatial resolution of 1 arc-second per pixel, (~ 30 meters per pixel at equator). Forest loss during the period 2000–2014, is defined as a stand-replacement disturbance, or a change from a forest to non-forest state, encoded as either 1 (loss) or 0 (no loss). Forest gain during the period 2000–2012, is defined as a non-forest to forest change entirely within the study period, encoded as either 1 (gain) or 0 (no gain).These layers were again cropped to the administrative boundaries of the Shimoga District (raster package: crop function) and re-projected to an equal area projection (Albers equal area conic projection) using the projectRaster function (method=”ngb”) in R. An algorithm was developed in R to identify which of the pixels in the loss and gain rasters coincided with each 1km and 2km grid cell of the study area. The ClassStat function of the SDM Tools package [4] was used to calculate the proportional area of each 1km or 2km grid cell that was made up of loss pixels or gain pixels. Forest gain and loss are very highly correlated (r=0.986) and occur in similar places in the landscape (Fig. S2C). Forest loss was a much more common transition than a forest gain affecting 1.2% of land pixels rather than 0.16% of land pixels for forest gain.

To assess how forest loss or gain from a global product like Hansen et al. (2013) should be interpreted locally in south India, we analysed how the loss and gain pixels from Hansen et al. 2013 coincided with classes in the MonkeyFeverRisk LULC map (by extracting the value of the LULC map for the centroids of loss or gain pixels).

The distribution of loss and gain pixels across forest classes from the MonkeyFeverRisk LULC map is shown in Table 1. Locations categorised as a loss by Hansen et al. were most commonly classified currently as plantation, followed by moist evergreen forest, followed by

moist or dry deciduous forest by the MonkeyFeverRisk LULC map. The pattern was similar for the gain pixels. Since not all forest loss pixels were non-forest in the current day and not all forest gain pixels were forest in the current day, the precise meaning of the Hansen et al. (2013) forest loss layer was unclear for south India, though we expect that it is at least indicative of areas where the forest has undergone a larger degree of change since 2000.

Table 1: Percentage of loss (n= 108398) and gain (n= 14646) land pixels from the global Hansen et al. (2013) product that fall into different forest classes according to the MonkeyFeverRisk LULC map

    Land use class


    Gain


    Loss




    moist evergreen


    30.4


    26.1




    moist deciduous


    6.5


    16.2




    dry deciduous


    3.0


    9.7




    plantation


    46.2


    37.2




    Non-forest classes


    14.0


    10.9

Host and public health predictors

Livestock host density data, namely buffalo and indigenous cattle densities in units of total head per village were obtained from Department of Animal Husbandry, Dairying and Fisheries, Government of India Census from 2011 at village level. These were linked to village boundaries from the Survey of India using the village census codes in R. The village areas were calculated from the spatial polygons dataframe of villages using the rgeos package in R, so that the total head per village metrics could be convert into an areal density of buffalo and indigenous cattle per km and then rasterized at 1km and 2km using the rasterize function of the raster package.

The human population size and public health metrics were obtained from the Government of India Population Census 2011. The human population size (census field TOT_P) was again linked to the spatial polygon village boundaries using the census village code (census field VCT_2011) and converted to an areal metric of population density per km and rasterized at 1km and 2km as above. The number of medics per head of population was derived by summing all doctors and para-medicals “in position” across all types of health centres, clinics and dispensaries per village and dividing by the total population of the village (TOT_P) and then linked to village boundaries and rasterized as above. The proximity to health centres was a categorical variable derived from the “Primary.Health.Centre..Numbers” field, where 1 = Primary Health Centre (PHC) within village boundary, 2 = PHC within 5km of village, 3=PHC within 5-10km of village, 4= PHC further than 10km from village. It was linked to village boundaries and rasterized as above.

The resulting raster layers for all predictors were saved in GeoTiff format.

References

Robert J. Hijmans (2017). raster: Geographic Data Analysis and Modeling. R package version 2.6-7. https://CRAN.R-project.org/package=raster
R Core Team (2018). R: A language and environment for statistical computing. R Foundation for Statistical Computing, Vienna, Austria.URL https://www.R-project.org/ 
Jarvis, A., Reuter, I., Nelson, A., Guevara, E. Hole-filled SRTM for the globe Version 4. 2008.
VanDerWal, J., Falconi, L., Januchowski, S., Shoo, L., & Storlie, C. (2014). SDMTools: Species Distribution Modelling Tools: Tools for processing data associated with species distribution modelling exercises. R

This map data product enriches traditional spatial coverage with real-time mobility data inputs. It provides a dynamic view of the built environment, shaped by actual human movement tracked from anonymized mobile device signals. Unlike static mapping layers, this map data evolves continuously to reflect changes in traffic flow, density, and behavioral zones.

The product includes: •Road network overlays with usage patterns •Venue and POI footprints enhanced by footfall counts •Transit corridors annotated with dwell and throughput data •Area-level mobility clusters and hotspots

Perfect for building routing applications, geo-visualizations, or spatial dashboards, this map data integrates seamlessly with location data, foot traffic data, and geospatial data infrastructures.

Use it to: •Identify underutilized or congested corridors•Prioritize capital investments or public service deployment •Enhance OOH advertising insights •Create visual simulations for urban design reviews

All data is provided in standard mapping formats (shapefiles, GeoJSON, raster tiles) and can be filtered by country, region, POI category, or date range.

This map data is continuously refined to ensure high fidelity, spatial granularity, and machine-readability. By tying static cartography to real-time behavior, it bridges the gap between what’s mapped and what’s happening

In the middle of 2023, about 60 percent of the global population was living in Asia.The total world population amounted to 8.1 billion people on the planet. In other words 4.7 billion people were living in Asia as of 2023. Global populationDue to medical advances, better living conditions and the increase of agricultural productivity, the world population increased rapidly over the past century, and is expected to continue to grow. After reaching eight billion in 2023, the global population is estimated to pass 10 billion by 2060. Africa expected to drive population increase Most of the future population increase is expected to happen in Africa. The countries with the highest population growth rate in 2024 were mostly African countries. While around 1.47 billion people live on the continent as of 2024, this is forecast to grow to 3.9 billion by 2100. This is underlined by the fact that most of the countries wit the highest population growth rate are found in Africa. The growing population, in combination with climate change, puts increasing pressure on the world's resources.

Values without references indicate that have been determined for this article (see S1 Fig).