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Literature studyLiterature considering the effect of pasture degradation on SOC, N, and clay content, as well as bulk density (BD), was assembled by searching (i) Web of Science V.5.22.1, (ii) ScienceDirect (Elsevier B.V.) (iii) Google Scholar, and (iv) the China Knowledge Resource Integrated Database (CNKI). Search terms were “degradation gradient”, “degradation stages”, “alpine meadow”, “Tibetan Plateau”, “soil”, “soil organic carbon”, and “soil organic matter” in different combinations. The criteria for including a study in the analysis were: (i) a clear and comprehensible classification of degradation stages was presented, (ii) data on SOC, N, and/or BD were reported, (iii) a non-degraded pasture site was included as a reference to enable an effect size analysis and the calculation of SOC and N losses, (iv) sampling depths and study location were clearly presented. (v) Studies were only considered that took samples in 10 cm depth intervals, to maintain comparability to the analyses from our own study site. The degradation stages in the literature studies were regrouped into the six successive stages (S0–S5) according to the respective degradation descriptions. In total, we compiled the results of 49 publications published between 2002 and 2020.When SOM content was presented, this was converted to SOC content using a conversion factor of 2.032. SOC and N stocks were calculated using the following equation:$${{{{{rm{Elemental; stock}}}}}}=100* {{{{{rm{content}}}}}}* {{{{{rm{BD}}}}}}* {{{{{rm{depth}}}}}}$$
(1)
where elemental stock is SOC or N stock [kg ha−1]; content is SOC or N content [g kg−1]; BD is soil bulk density [g cm−3] and depth is the soil sampling depth [cm].The effect sizes of individual variables (i.e., SOC and N stocks as well as BD) were quantified as follows:$${{{{{rm{ES}}}}}}=,frac{(D-R)}{R* 100 % }$$
(2)
where ES is the effect size in %, D is the value of the corresponding variable in the relevant degradation stage and R is the value of each variable in the non-degraded stage (reference site). When ES is positive, zero, or negative, this indicates an increase, no change, or decrease, respectively, of the parameter compared to the non-degraded stage.Experimental design of the field studyLarge areas in the study region are impacted by grassland degradation. In total, 45% of the surface area of the Kobresia pasture ecosystem on the TP is already degraded2. The experiment was designed to differentiate and quantify SOC losses by erosion vs. net decomposition and identify underlying shifts in microbial community composition and link these to changes in key microbial functions in the soil C cycle. We categorized the range of Kobresia root-mat degradation from non-degraded to bare soils into six successive degradation stages (S0–S5). Stage S0 represented non-degraded root mats, while stages S1–S4 represented increasing degrees of surface cracks, and bare soil patches without root mats defined stage S5 (Supplementary Fig. 1). All six degradation stages were selected within an area of about 4 ha to ensure equal environmental conditions and each stage was sampled in four field replicates. However, the studied degradation patterns are common for the entire Kobresia ecosystem (Supplementary Fig. 1).Site descriptionThe field study was conducted near Nagqu (Tibet, China) in the late summer 2013 and 2015. The study site of about 4 ha (NW: 31.274748°N, 92.108963°E; NE: 31.274995°N, 92.111482°E; SW: 31.273488°N, 92.108906°E; SE: 31.273421°N, 92.112025°E) was located on gentle slopes (2–5%) at 4,484 m a.s.l. in the core area of the Kobresia pygmaea ecosystem according to Miehe et al.8. The vegetation consists mainly of K. pygmaea, which covers up to 61% of the surface. Other grasses, sedges, or dwarf rosette plants (Carex ivanoviae, Carex spp., Festuca spp., Kobresia pusilla, Poa spp., Stipa purpurea, Trisetum spp.) rarely cover more than 40%. The growing season is strongly restricted by temperature and water availability. At most, it lasts from mid-May to mid-September, but varies strongly depending on the onset and duration of the summer monsoon. Mean annual precipitation is 431 mm, with roughly 80% falling as summer rains. The mean annual temperature is −1.2 °C, while the mean maximum temperature of the warmest month (July) is +9.0 °C2.A characteristic feature of Kobresia pastures is their very compact root mats, with an average thickness of 15 cm at the study site. These consist mainly of living and dead K. pygmaea roots and rhizomes, leaf bases, large amounts of plant residue, and mineral particles. Intact soil is a Stagnic Eutric Cambisol (Humic), developed on a loess layer overlying glacial sediments and containing 50% sand, 33% silt, and 17% clay in the topsoil (0–25 cm). The topsoil is free of carbonates and is of neutral pH (pH in H2O: 6.8)5. Total soil depth was on average 35 cm.The site is used as a winter pasture for yaks, sheep, and goats from January to April. Besides livestock, large numbers of plateau pikas (Ochotona) are found on the sites. These animals have a considerable impact on the plant cover through their burrowing activity, in particular the soil thrown out of their burrows, which can cover and destroy the Kobresia turf.Sampling designThe vertical and horizontal extent of the surface cracks was measured for each plot (Supplementary Table 2). Vegetation cover was measured and the aboveground biomass was collected in the cracks (Supplementary Table 2). In general, intact Kobresia turf (S0) provided high resistance to penetration as measured by a penetrologger (Eijkelkamp Soil and Water, Giesbeek, NL) in 1 cm increments and four replicates per plot.Soil sampling was conducted using soil pits (30 cm length × 30 cm width × 40 cm depth). Horizons were classified and then soil and roots were sampled for each horizon directly below the cracks. Bulk density and root biomass were determined in undisturbed soil samples, using soil cores (10 cm height and 10 cm diameter). Living roots were separated from dead roots and root debris by their bright color and soft texture using tweezers under magnification, and the roots were subsequently washed with distilled water to remove the remaining soil. Because over 95% of the roots occurred in the upper 25 cm5, we did not sample for root biomass below this depth.Additional soil samples were taken from each horizon for further analysis. Microbial community and functional characterization were performed on samples from the same pits but with a fixed depth classification (0–5 cm, 5–15 cm, 15–35 cm) to reduce the number of samples.Plant and soil analysesSoil and roots were separated by sieving (2 mm) and the roots subsequently washed with distilled water. Bulk density and root density were determined by dividing the dry soil mass (dried at 105 °C for 24 h) and the dry root biomass (60 °C) by the volume of the sampling core. To reflect the root biomass, root density was expressed per soil volume (mg cm−3). Soil and root samples were milled for subsequent analysis.Elemental concentrations and SOC characteristicsTotal SOC and total N contents and stable isotope signatures (δ13C and δ15N) were analyzed using an isotope ratio mass spectrometer (Delta plus, Conflo III, Thermo Electron Cooperation, Bremen, Germany) coupled to an elemental analyzer (NA 1500, Fisons Instruments, Milano, Italy). Measurements were conducted at the Centre for Stable Isotope Research and Analysis (KOSI) of the University of Göttingen. The δ13C and δ15N values were calculated by relating the isotope ratio of each sample (Rsample = 13C/12C or 15N/14N) to the international standards (Pee Dee Belemnite 13C/12C ratio for δ13C; the atmospheric 15N/14N composition for δ15N).Soil pH of air-dried soil was measured potentiometrically at a ratio (v/v) of 1.0:2.5 in distilled water.Lignin phenols were depolymerized using the CuO oxidation method25 and analyzed with a gas chromatography-mass spectrometry (GC–MS) system (GC 7820 A, MS 5977B, Agilent Technologies, Waldbronn, Germany). Vanillyl and syringyl units were calculated from the corresponding aldehydes, ketones, and carboxylic acids. Cinnamyl units were derived from the sum of p-coumaric acid and ferulic acid. The sum of the three structural units (VSC = V + S + C) was considered to reflect the lignin phenol content in a sample.DNA extraction and PCRSamples were directly frozen on site at −20 °C and transported to Germany for analysis of microbial community structure. Total DNA was extracted from the soil samples with the PowerSoil DNA isolation kit (MoBio Laboratories Inc., Carlsbad, CA, USA) according to the manufacturer’s instructions, and DNA concentration was determined using a NanoDrop 1000 spectrophotometer (Thermo Fisher Scientific, Wilmington, DE, USA). The extracted DNA was amplified with forward and reverse primer sets suitable for either t-RFLP (fluorescence marked, FAM) or Illumina MiSeq sequencing (Illumina Inc., San Diego, USA): V3 (5’-CCT ACG GGN GGC WGC AG-3’) and V4 (5’-GAC TAC HVG GGT ATC TAA TCC-3’) primers were used for bacterial 16 S rRNA genes whereas ITS1 (5’-CTT GGT CAT TTA GAG GAA GTA A-3’), ITS1-F_KYO1 (5’-CTH GGT CAT TTA GAG GAA STA A-3’), ITS2 (5’-GCT GCG TTC TTC ATC GAT GC-3’) and ITS4 (5’-TCC TCC GCT TAT TGA TAT GC-3’) were used for fungi33,34. Primers for Illumina MiSeq sequencing included adaptor sequences (forward: 5’-TCG TCG GCA GCG TCA GAT GTG TAT AAG AGA CAG-3’; reverse: 5’-GTC TCG TGG GCT CGG AGA TGT GTA TAA GAG ACA G-3’)33. PCR was performed with the Phusion High-Fidelity PCR kit (New England Biolabs Inc., Ipswich, MA, USA) creating a 50 µl master mix with 28.8 µl H2Omolec, 2.5 µl DMSO, 10 µl Phusion GC buffer, 1 µl of forward and reverse primer, 0.2 µl MgCl2, 1 µl dNTPs, 0.5 µl Phusion HF DNA Polymerase, and 5 µl template DNA. PCR temperatures started with initial denaturation at 98 °C for 1 min, followed by denaturation (98 °C, 45 s), annealing (48/60 °C, 45 s), and extension (72 °C, 30 s). These steps were repeated 25 times, finalized again with a final extension (72 °C, 5 min), and cooling to 10 °C. Agarose gel electrophoresis was used to assess the success of the PCR and the amount of amplified DNA (0.8% gel:1.0 g Rotigarose, 5 µl Roti-Safe Gelstain, Carl Roth GmbH & Co. KG, Karlsruhe, Germany; and 100 ml 1× TAE-buffer). PCR product was purified after initial PCR and restriction digestion (t-RFLP) with either NucleoMag 96 PCR (16 S rRNA gene amplicons, Macherey-Nagel GmbH & Co. KG, Düren, Germany) or a modified clean-up protocol after Moreau (t-RFLP)35: 3× the volume of the reaction solution as 100% ethanol and ¼x vol. 125 mM EDTA was added and mixed by inversion or vortex. After incubation at room temperature for 15 min, the product was centrifuged at 25,000 × g for 30 min at 4 °C. Afterwards the supernatant was removed, and the inverted 96-well plate was centrifuged shortly for 2 min. Seventy microliters ethanol (70%) were added and centrifuged at 25,000 × g for 30 min at 4 °C. Again, the supernatant was removed, and the pallet was dried at room temperature for 30 min. Finally, the ethanol-free pallet was resuspended in H2Omolec.T-RFLP fingerprintingThe purified fluorescence-labeled PCR products were digested with three different restriction enzymes (MspI and BstUI, HaeIII) according to the manufacturer’s guidelines (New England Biolabs Inc., Ipswich, MA, USA) with a 20 µl master mix: 16.75 µl H2Omolec, 2 µl CutSmart buffer, 0.25 or 0.5 µl restriction enzyme, and 1 µl PCR product for 15 min at 37 °C (MspI) and 60 °C (BstUI, HaeIII), respectively. The digested PCR product was purified a second time35, dissolved in Super-DI Formamide (MCLAB, San Francisco, CA, USA) and, along with Red DNA size standard (MCLAB, San Francisco, USA), analyzed in an ABI Prism 3130 Genetic Analyzer (Applied Biosystems, Carlsbad, CA, USA). Terminal restriction fragments shorter than 50 bp and longer than 800 bp were removed from the t-RFLP fingerprints.16 S rRNA gene and internal transcribed spacer (ITS) sequencing and sequence processingThe 16 S rRNA gene and ITS paired-end raw reads for the bacterial and fungal community analyses were deposited in the National Center for Biotechnology Information (NCBI) Sequence Read Archive (SRA) and can be found under the BioProject accession number PRJNA626504. This BioProject contains 70 samples and 139 SRA experiments (SRR11570615–SRR11570753) which were processed using CASAVA software (Illumina, San Diego, CA, USA) for demultiplexing of MiSeq raw sequences (2 × 300 bp, MiSeq Reagent Kit v3).Paired-end sequences were quality-filtered with fastp (version 0.19.4)36 using default settings with the addition of an increased per base phred score of 20, base-pair corrections by overlap (-c), as well as 5′- and 3′-end read trimming with a sliding window of 4, a mean quality of 20 and minimum sequence size of 50 bp. Paired-end sequences were merged using PEAR v0.9.1137 with default parameters. Subsequently, unclipped reverse and forward primer sequences were removed with cutadapt v1.1838 with default settings. Sequences were then processed using VSEARCH (v2.9.1)39. This included sorting and size-filtering (—sortbylength,—minseqlength) of the paired reads to ≥300 bp for bacteria and ≥140 bp for ITS1, dereplication (—derep_fulllength). Dereplicated sequences were denoised with UNOISE340 using default settings (—cluster_unoise—minsize 8) and chimeras were removed (—uchime3_denovo). An additional reference-based chimera removal was performed (—uchime_ref) against the SILVA41 SSU NR database (v132) and UNITE42 database (v7.2) resulting in the final set of amplicon sequence variants (ASVs)43. Quality-filtered and merged reads were mapped to ASVs (—usearch_global–id 0.97). Classification of ASVs was performed with BLAST 2.7.1+ against the SILVA SSU NR (v132) and UNITE (v7.2) database with an identity of at least 90%. The ITS sequences contained unidentified fungal ASVs after UNITE classification, these sequences were checked (blastn)44 against the “nt” database (Nov 2018) to remove non-fungal ASVs and only as fungi classified reads were kept. Sample comparisons were performed at the same surveying effort, utilizing the lowest number of sequences by random selection (total 15,800 bacteria, 20,500 fungi). Species richness, alpha and beta diversity estimates, and rarefaction curves were determined using the QIIME 1.9.145 script alpha_rarefaction.py.The final ASV tables were used to compute heatmaps showing the effect of degradation on the community using R (Version 3.6.1, R Foundation for Statistical Computing, Vienna, Austria) and R packages “gplots”, “vegan”, “permute” and “RColorBrewer”. Fungal community functions were obtained from the FunGuild database46. Plant mycorrhizal association types were compiled from the literature38,39,40,41,47,48,49,50. If no direct species match was available, the mycorrhizal association was assumed to remain constant within the same genus.Enzyme activityEnzyme activity was measured to characterize the functional activity of the soil microorganisms. The following extracellular enzymes, involved in C, N, and P transformations, were considered: two hydrolases (β-glucosidase and xylanase), phenoloxidase, urease, and alkaline phosphatase. Enzyme activities were measured directly at the sampling site according to protocols after Schinner et al.51. Beta-glucosidase was incubated with saligenin for 3 h at 37 °C, xylanase with glucose for 24 h at 50 °C, phenoloxidase with L-3,4-dihydroxy phenylalanine (DOPA) for 1 h at 25 °C, urease with urea for 2 h at 37 °C and alkaline phosphatase on P-nitrophenyl phosphate for 1 h at 37 °C. Reaction products were measured photometrically at recommended wavelengths (578, 690, 475, 660, and 400 nm, respectively).SOC stocks and SOC lossThe SOC stocks (in kg C m−2) for the upper 30 cm were determined by multiplying the SOC content (g C kg−1) by the BD (g cm−3) and the thickness of the soil horizons (m). SOC losses (%) were calculated for each degradation stage and horizon and were related to the mean C stock of the reference stage (S0). The erosion-induced SOC loss of the upper horizon was estimated by considering the topsoil removal (extent of vertical soil cracks) of all degraded soil profiles (S1–S5) and the SOC content and BD of the reference (S0). To calculate the mineralization-derived SOC loss, we accounted for the effects of SOC and root mineralization on both SOC content and BD. Thus, we used the SOC content and BD from each degradation stage (S1–S5) and multiplied it by the mean thickness of each horizon (down to 30 cm) from the reference site (S0). The disentanglement of erosion-derived SOC loss from mineralization-derived SOC loss was based on explicit assumptions that (i) erosion-derived SOC losses are mainly associated with losses from the topsoil, and (ii) the decreasing SOC contents in the erosion-unaffected horizons were mainly driven by mineralization and decreasing root C input.Statistical analysesStatistical analyses were performed using PASW Statistics (IBM SPSS Statistics) and R software (Version 3.6.1). Soil and plant characteristics are presented as means and standard errors (means ± SE). The significance of treatment effects (S0–S5) and depth was tested by one-way ANOVA at p  More

Study siteWe conducted this research at the Uluwatu temple site in Bali, Indonesia. Uluwatu is located on the Island’s southern coast, in the Badung Regency. The temple at Uluwatu is a Pura Luhur, which is a significant temple for Balinese Hindus across the island and is therefore visited regularly for significant regional, community, family, and household rituals by Balinese people from different regions throughout the year18. During the period of data collection hundreds of tourists also visit the Uluwatu temple each day. The temple sits on top of a promontory cliff edge, with walking paths in front of it that continue in loops to the North and South. These looping pathways surround scrub forests, which the macaques frequently inhabit but the humans rarely enter.In 2017–2018 there were five macaque groups at Uluwatu, which ranged throughout the temple complex area, and beyond. All groups are provisioned daily with a mixed diet of corn, cucumbers, and bananas by temple staff members. The two groups included in this research are the Celagi and Riting groups. We selected these groups because they previously exhibited significant differences in robbing frequencies whereby Riting was observed exhibiting robbing and bartering more frequently than Celagi1. Furthermore, both groups include the same highly trafficked tourist areas in their overlapping home ranges relative to the other groups at Uluwatu, theoretically minimizing between group differences in the contexts of human interaction1,19.Data collectionJVP collected data from May, 2017 to March, 2018 totaling 197 focal observation hours on all 13 subadult males in Celagi and Riting that were identified in May–June 2017. Subadult male long-tailed macaques exhibit characteristic patterns of incomplete canine eruption, sex organ development, and body size growth, which achieves a maximum of 80% of total adult size18. Mean sampling effort per individual was 15.2 hours (h), with a range of 1.75 h, totaling 102.75 h for Riting and 94.75 h for Celagi. The data collection protocol consisted of focal-animal sampling and instantaneous scan sampling20 on all six subadult males in the Celagi group, and all seven subadult males in the Riting group. Focal follows were 15 minutes in length. Sampling effort per individual is presented in Table 1. A random number generator determined the order of focal follows each morning. In the event a target focal animal could not be located within 10 minutes of locating the group, the next in line was located and observed. Data presented here come from focal animal sampling records of state and event behaviors. Relevant event behaviors consist of agonistic gestures used for calculating dominance relationships, including the target, or interaction partner, of all communicative event behaviors and the time of its occurrence. All changes in the focal animal’s state behavior were noted, recording the time of the change to the minute.Table 1 Focal Subadult male long-tailed macaques in Celagi and Riting at Uluwatu, Bali, Indonesia.Full size tableDuring focal samples we recorded robbing and bartering as a sequence of mixed event and state behaviors. We scored both the robbery and exchange phases as event behaviors, and the interim phase of item possession as a state behavior. We record a robbery as successful if the focal animal took an object from a human and established control of the object with their hands or teeth, and as unsuccessful if the focal animal touched the object but was not able to establish control of it. For each successful robbery we recorded the object taken. Unsuccessful robberies end the sequence, whereas successful robberies are typically followed by various forms of manipulating the object.The robbing and bartering sequence ends with one of several event behavior exchange outcomes: (1) “Successful exchanges” consist of the focal animal receiving a food reward from a human and releasing the stolen object; (2) “forced exchanges” are when a human takes the object back without a bartering event; (3) “dropped objects” describe when the macaque loses control of the object while carrying it or otherwise locomoting, and is akin to an “accidental drop”; (4) “no exchange” includes instances of the macaque releasing the object for no reward after manipulating it; and (5) “expired observation” consists of instances in which the final result of the robbing and bartering event was unobserved in the sample period (i.e., the sample period ended while the macaque still had possession of the object). A 6th exchange outcome is “rejected exchange,” which occurs when the focal animal does not drop the stolen object after being offered, or in some cases even accepting, a food reward. The “rejected exchange” outcome is unique in that it does not end the robbing and bartering sequence because a human may have one or more exchange attempts rejected before eventually facilitating a successful exchange, or before one of the other outcomes (2–5) occurs. For each successful exchange we recorded the food item the macaques received. Food items are grouped into four categories: fruits, peanuts, eggs, and human snacks. Snacks include packaged and processed food items such as candy or chips.Data analysisWe grouped the broad range of stolen items into classes of general types. “Eyewear” combines eyeglasses and sunglasses, while “footwear” combines sandals and shoes. “Ornaments” includes objects attached to and/or hanging from backpacks, such as keychains, while “accessories” includes decorative objects attached to an individual’s body or clothing like bracelets and hair ties. “Electronics” covers cellular phones and tablets. “Hats” encompasses removable forms of headwear, most typically represented by baseball-style hats or sun hats. “Plastics” is an item class consisting of lighters and bottles, which may be filled with water, soda, or juice. The “unidentified” category is used for stolen items which could not be clearly observed during or after the robbing and bartering sequence.“Robbery attempts” refers to the combined total number of successful and unsuccessful robberies. “Robbery efficiency” is a novel metric referring to the number of successful robberies divided by the total number of robbery attempts. The “Exchange Outcome Index” is calculated by dividing the number of successful exchanges by the total number of robbery attempts. We make this calculation using robbery attempts instead of successful robberies to account for total robbery effort because failed robberies still factor into an individual’s total energy expenditure toward receiving a bartered food reward and their total exposure to the risks (e.g., physical retaliation) of stealing from humans relative to achieving the desired end result of a food reward.Social rank was measured with David’s Score, calculated using dyadic agonistic interactions. We coded “winners” of contests as those who exhibited the agonistic behavior, while “losers” were the recipients of those agonistic behaviors21,22. We excluded intergroup agonistic interactions in our calculations of David’s Score.To account for potential variation in the overall patterns of interaction with humans between groups we calculated a Human Interaction Rate, which is the sum of human-directed interactions from focal animals in each group divided by the total number of observation hours on focal animals in that group.Statistical analysisWe ran statistical tests in SYSTAT software with a significance level set at 0.05. We used chi-square goodness-of-fit tests to assess the significance of differences in successful robberies between individuals for each group. To avoid having cells with values of zero, two focal subjects, Minion and Spot from Celagi, are excluded from this test because neither were observed making a successful robbery during the observation period. We also used chi-square goodness-of-fit tests to assess exchange outcome occurrences within each group, as well as a Fisher’s exact to test for significant differences in robbery outcomes between groups due to low expected counts in 40% of the cells. “Rejected exchange” events were not included in the analysis of robbery outcomes because they do not end the sequence and are therefore not mutually exclusive with the other robbery outcomes.We further tested for the effect of dominance position on robbery outcomes. Due to our small sample size and the preliminary nature of this investigation, we used Spearman correlations to assess the relationship between subadult male dominance position via David’s Score and (1) robbing efficiency and (2) the Exchange Outcome Index.Compliance with ethical standardsThis research complied with the standards and protocols for observational fieldwork with nonhuman primates and was approved by the University of Notre Dame Compliance IACUC board (protocol ID: 16-02-2932), where JVP and AF were affiliated at the time of this research. This study did not involve human subjects. This research further received a research permit from RISTEK in Indonesia (permit number: 2C21EB0881-R), and complied with local laws and customary practices in Bali. More

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Growth in harvests of crops meant for exports, processing and industrial use, together with their higher yields and faster yield gains, stands out globally; at a more granular level, this was driven by specific global regions that are getting increasingly specialized in harvesting crops for these usages.Changes in global-level harvested areasAt the global scale, we find that crops harvested for direct food utilization have the highest area and have been relatively stable over the study period (Fig. 1a). However, as the total harvested hectares have increased globally (Supplementary Table 1), this has translated into decreasing fractions of crops harvested for direct food utilization, from ~51% in the 1960s (average over 1964 to 1968) to ~37% in the 2010s (average over 2009 to 2013), with a similar reduction in feed crop harvests (Table 1). Conversely, there has been a substantial increase in crops for processing, exports and industrial use (Fig. 1a, Table 1 and Supplementary Table 1). The increase in industrial crop harvests occurred after year 2000. Around the same time, harvested hectares for exported crops ramped up and by the 2010s had surpassed those of crops harvested for feed use (Fig. 1a). Crops harvested for seed usage and losses are relatively minor, and we will not discuss them further. If the global trends observed in the past 20 years continue (Fig. 1a), by 2030, crops harvested for exports, processing and industrial use will account for ~ 23%, 17% and 8% of overall harvested hectares, whereas those for food will decrease to ~29% (Table 1).Fig. 1: Sector-based global crop-utilization trends.a–d, Observed total harvested ha (a), average yield in kcal ha−1 per year (b), average yield in protein ha−1 per year (c) and average yield in fat ha−1 per year (d) in the seven sectors of food, feed, processing, export, other uses (non-food/industrial), seed and losses from 1964 to 2013, annually, and projections to 2030 based on the past 20 years. The shading shows the 90% confidence interval for the significant linear model projections.Full size imageTable 1 Sector-based global crop-utilization changesFull size tableChanges in global-level crop yieldsWe find that crops harvested for direct food usage generally have had lower yields than all other sectors at the global scale over the time period of the study (Fig. 1b–d). This is not a new phenomenon, as crops harvested for direct food utilization have always had lower yields relative to other sectors (Supplementary Table 1). What has changed, however, is the ramping up (steeper positive slopes) of industrial, export and processing crop yields (Fig. 1b–d and Table 1). At these rates, caloric yields of industrial-use crops could increase by 28% from the 2010s to 2030 compared with 24% and 21% yield increases of crops harvested for directly consumed food and for feed use (Fig. 1b). Given that caloric yields of industrial-use crops are already substantially higher than food and feed crops (2× and 1.4×, respectively, in the 2010s), the faster caloric yield increases for industrial-use crops will widen this gap (2.1× and 1.5×, respectively). Yield measurements in other units of protein and fat show similar results (Table 1, Fig. 1c,d and Supplementary Table 1).Changes in the spatial patterns of harvested areas and productionWithin country-level information on harvested areas and productivity based on utilization categories is required for developing more locally effective agricultural policies. Over the course of the study time period 1964 to 2013 (Fig. 2a,b and Supplementary Video 1), we find changes in all continents when spatially analysed at the grid-cell level, except for most parts of Africa. Even in Africa, there are locations with fractional reductions in food crop harvests over the study period, such as parts of Angola, Ghana, Nigeria and South Africa. Within these and other countries, the exact location, magnitude and direction of the change varies from one region to the next (that is, compare Fig. 2a with Fig. 2b).Fig. 2: Sector-based spatial changes in crop harvests.a,b, The fraction of a grid cell in one of seven categories—food, feed, processing, export, other (non-food/industrial use), seed and losses—in each period, 1964–1968 (a) and 2009–2013 (b).Full size imageCrops harvested for direct food utilization have been prevalent in Asia, though much has changed since the 1960s (Fig. 2a,b and Supplementary Video 1). In China, there appears to be an imaginary belt, north and west of which harvests of crops used as directly consumed food decreased between the 1960s (Fig. 2a) and 2010s (Fig. 2b), while those for other uses increased. This belt appears to roughly extend from the northern half of Jiangsu (a province on the Yellow Sea in the east), curving westwards and southwards through northern Anhui, southern Henan, central Hubei and the northern tip of Hunan, and then turning sharply south and splitting Guangdong (a province on the South China Sea) through the middle. The sector gaining from the 10–20% fractional food harvest reduction varies. The increase in crops for feed, processing and industrial usage increases as one moves northward, especially north of Jiangsu and Anhui (Fig. 2a,b and Supplementary Video 1).Similarly, in India, there is a north–south zone encompassing eastern Haryana in the north, moving southwards through eastern Rajasthan, western Madhya Pradesh to eastern Maharashtra in the south, where there was a drastic reduction in crops harvested for direct food utilization over the study period (Fig. 2 and Supplementary Video 1); crops harvested for processing primarily increased. Changes in South and Southeast Asia over the study period are primarily away from once-dominant harvests of directly consumed food crops to feed crops, followed by processing crops, export crops and industrial-use crops, as in Myanmar and Thailand. In Malaysia, the growth was in export and industrial-usage crops, whereas in Indonesia, it was export crops and smaller increases in industrial-utilization crops. Central Asian states, especially Kazakhstan and some parts of Russia, witnessed a large reduction in crops harvested for direct food use over the study period, replaced by the crops destined for exports between the two periods (Fig. 2 and Supplementary Video 1).In Australia in the 1960s, food crops were harvested everywhere, accounting for ~10% of the total, which declined to ~5% by the 2010s. This was accompanied by small reductions in crops harvested for feed and export and balanced mainly by increases in crops for processing and industrial utilization (Fig. 2 and Supplementary Video 1).In Europe in the 1960s, crops were dominantly harvested for food and feed, but by the 2010s, this changed to include crops harvested for processing (Fig. 2 and Supplementary Video 1). In France, major reductions in feed crops have been balanced by growth in processing, export and industrial-use crops. In Spain, the primary change is from crops harvested for direct food to those of feed. In Germany, crops harvested for export have replaced those for direct food utilization.Latin America used to dominantly harvest food crops (as in Mexico) or food and feed crops (as in Brazil and Argentina) (Fig. 2 and Supplementary Video 1). Midwestern Brazil used to harvest only food crops, and feed and processing crop harvests were restricted to the Atlantic states (the 1960s; Fig. 2a), but by the 2010s (Fig. 2b), harvests of food crops had become a negligible fraction in Midwestern Brazil (as in Mato Grosso), and crops harvested for processing and exports are dominant now. In the Atlantic states of Brazil, one of the major changes is the increased proportion of harvests for industrial crops. In Argentina, over the study period, the proportion of crops harvested for food and feed has decreased, and this utilization has been mainly replaced by crops harvested for processing; crops harvested for exports changed, but the direction of change was spatially heterogeneous across Argentina (Fig. 2 and Supplementary Video 1). In Mexico, the primary change is the reduction in the fraction of crops harvested for direct food consumption and the increased harvests of crops for feed.Crops harvested for food and feed are also on the decline proportionally in North America. The United States has experienced a change from the dominance of food and feed crops in the 1960s to processing and industrial-usage crops in the 2010s. Detailed changes in the United States and Canada vary from one location to the next (Fig. 2), though the major change is the lower fraction of crops harvested for direct food consumption.Results are similar when viewed through the lens of calories, protein and fat with local-level differences as yields vary based on the measurement units (Supplementary Fig. 1). Further dramatic changes can be expected if observed linear trends from 1994 to 2013 at each grid cell continued until 2030 (Supplementary Fig. 2).Calories harvested in 2030 and achieving UN SDG 2We compare the extra food calories that will potentially be harvested in 2030 (Fig. 3a and Supplementary Data 2) to those required for both the projected extra population and feeding the projected undernourished population in each country (Fig. 3b and Supplementary Data 2). As an extreme case, we also compared whether total calories (all seven utilization sectors) would be sufficient (Fig. 3c and Supplementary Data 2). Altogether, we evaluated 156 countries, of which 86 had reported undernourished populations (Supplementary Data 2). On the basis of the minimum dietary energy requirement (MDER), we find that countries with reported undernourished populations will have a shortfall of ~675.4 trillion kcal per year to nourish the increased population and the expected undernourished from their extra harvested food calories. However, compared with the more realistic average dietary energy requirement (ADER), this shortfall will be ~993.9 trillion kcal per year (or ~70% from requirements) in 2030 (15 additional scenarios of undernourished populations in 2030 (provided in Supplementary Data 3) show global calorie shortfalls may similarly range from ~587.2 trillion kcal per year to ~1,269.3 trillion kcal per year based on the MDER level of nutrition requirement, and ~880.7 trillion kcal per year to ~1,755.6 trillion kcal per year based on the more realistic ADER level of nutrition requirement in 2030).Fig. 3: Meeting UN SDG goal 2 in 2030.a, Same as Fig. 2 but for the projected kcal ha−1 per year in 2030 per utilization sector and then mapping the fraction of total kcal ha−1 per year projected as harvested. b, Shortfall or gap from kcal per year harvested in 2030 as crops for direct food use and those to plug the gap from population growth and/or undernourished population. Computed based on the 2018 to 2020 ADER number for the country. c, Same as b but the kcal per year harvested used for computation is the total across all the seven sectors and shortfall is from whether the total calories harvested were used for direct food consumption (little to no processing).Full size imageCountries reporting undernourishment can, however, meet their requirement of extra calories in 2030 for both population change and those for the undernourished if calories from other utilization sectors are diverted and consumed directly as food calories (Fig. 3c and Supplementary Data 2 and 3). Though at the global scale, it appears that countries with high levels of undernourishment in 2030 can divert just a portion of their total harvested calories and meet some of the requirements of UN’s SDG 2 (ref. 4). In reality, many of the individual countries concentrated in sub-Saharan Africa have limited scope of diversion of calories from other sectors such as feed, processing or exports as crops for direct food use, as they already harvest most crops for direct food consumption (Fig. 2 and Supplementary Figs. 1 and 2). As such, many countries in this region may see deepening reliance on food imports. Note that the UN’s second SDG goal is broader in scope, including efforts to end malnutrition and increase agricultural productivity, among other goals4. Reconfiguration planning19 can use our spatially detailed information (Figs. 2 and 3, Supplementary Figs. 1 and 2 and Supplementary Data 2 and 3) in conjunction with policies that incentivize increased food crop harvests globally and ensure their equitable distribution to undernourished regions when local production is not sufficient20,21. This will require supply chain management22,23 and detailed analysis of optimization scenarios24 with our maps and tables as an important step linking specific production regions with the initial use of that production. More