Philippa Kaur

Disturbance detectionWe used a well-established spectral-temporal segmentation method, Landsat-based Detection of Trends in Disturbance and Recovery (LandTrendr), to detect disturbances within the Google Earth Engine (GEE) cloud-computing platform57,58. The core of the LandTrendr is to extract a set of disturbance-related metrics by breaking pixel-level annual time-series spectral trajectories into linear features using Landsat observations. The LandTrendr has been widely used for change detection in various forest settings, and details about the algorithms can be found in previous publications57. Here we briefly described the key steps in generating the year and type of disturbances in China’s forests using the LandTrendr within the GEE platform. The overall analytic flow can be found in Supplementary Fig. 10.First, we generated annual spectrally consistent time-series data by using all available, good quality (cloud cover ≤ 20) Tier 1 Landsat 5 (Thematic Mapper), Landsat 7 (Enhanced Thematic Mapper Plus), and Landsat 8 (Operational Land Imager) images acquired during the peak growing seasons (June 1—September 30) from 1986 to 2020. The peak growing seasons were selected to exclude compounding influences from ice, snow, and soil, and to maximize the spectral changes after forest disturbances. To tackle the spectral inconsistency among Landsat sensors, we harmonized spectral values via linear transformations according to band-respective coefficients presented in59. Clouds, cloud shadows, snow, and water were masked out using the Fmask algorithm60. The annual band composites at 30-meter spatial resolution during 1986–2020 were computed using the Medoid method61.Secondly, we ran the LandTrendr using five spectral indices, including two spectral bands (shortwave infrared I and II that were B5 and B7), tasseled cap wetness (TCW), normalized burn ratio (NBR), and normalized difference vegetation index. These five indices were effective indictors to represent vegetation greenness and structures, and were commonly used for detecting changes in forest disturbance and recovery62. For each spectral index, the LandTrendr produced a set of parameters to describe a possible disturbance event at the pixel level, including spectral values at pre-disturbance level (preval), magnitude of change (mag), duration (dur) and rate of change (rate), and the signal-to-noise ratio (dsnr) (n = 5). Using these five spectral indices, we generated a stack of disturbance-related parameter layers (n = 25, 5 spectral indices × 5 parameters), which were later used to detect and classify disturbances using machine learning models derived from reference data (described below).Disturbance classificationReference dataHigh-quality consistent reference data is key to train and classify disturbance types. To do so, we generated a total of 31225 reference points using a hierarchical approach. We first generated a large number of potential disturbance points using forest loss data from 2001 to 20203. Then we separated fire disturbances from non-fire disturbances by overlaying MODIS burned area (BA) with potential disturbance points following the procedure used by63. Specifically, fire disturbances were determined if the MODIS BA data coincided with the Landsat-derived forest loss for the fire year and 2 years postfire (i.e., t + 0, t + 1, t + 2) to account for delayed post-fire tree mortality. Following this step, we derived points as potential disturbances that consisted of fires and non-fire disturbances (including forest conversion to other land use types and silvicultural practices at various intensities). We also generated roughly the same number of points that experienced no disturbances (e.g., persistent forests), which were determined by selecting pixels with very few changes in spectral indices. These reference points, including fire, non-fire disturbances, and persistent forests, were then used to sample the time-series spectral data from 1986 to 2020. Finally, time-series spectral data from each reference point were visually checked to make sure they accurately represented disturbance events. This process resulted into a total of 31225 reference data points, including 2356 fire disturbance points, 13,242 non-fire disturbance points, and 15,627 no disturbance points (persistent forests) (Supplementary Fig. 2).Random forest classificationWe used machine learning modeling to classify each pixel into fire disturbance, non-fire disturbance, or no disturbance. The reference data points were used to sample the LandTrendr-derived disturbance-related parameter layers described above, which resulted into a dataset consisting of disturbance types. We divided the dataset into 70% of training data, and 30% as validation data. Using the training data, a Random Forest (RF) model was trained to classify each reference point into fire, non-fire disturbance, or no disturbance. Our RF approach showed that short-wave infrared (SWIR)-based moisture indices (e.g., B7, TCW) were strong predictors for detecting forest disturbances (Supplementary Fig. 11) likely because of their sensitivity to vegetation water content and canopy structure64. Finally, we applied the trained RF model to the full classification stack to consistently map the disturbance types from 1986 to 2020 across China’s forests, assuming that the spectral trajectories derived from reference data period 2001–2020 can be extrapolated to the whole mapping period 1986–2020. However, note that our approach was meant to detect relatively acuate and discrete disturbances that caused canopy opening, rather than subtle changes of forest structure or composition resulted from low intensive silvicultural practices and chronic disturbances.Year of disturbanceWe used the LandTrendr to determine the year of disturbance as the onset of magnitude of spectral change. Since we ran LandTrendr on five spectral indices, there were five possible years of disturbance for each pixel. Thus, we determined the year of disturbance using the median value from at least three different indices (i.e., NDVI, NBR, TCW, B5, B7). In this way, we only kept pixels that were detected as disturbances using at least three indices, thus reducing commission errors. The year with the greatest spectral changes generated by the LandTrendr often had an accuracy within 3 years11. A confidence level was also assigned to each disturbed pixel based on numbers of indices which showed possible disturbance events. Specially, low, medium, and high confidence were assigned if the disturbance was detected by three, four, or five spectral indices, respectively.ValidationsWe validated the disturbance map at the pixel and national levels. At the pixel level, we validated the final map using the validation sub-sample described in the previous section. We derived a confusion matrix to report user’s and producer’s accuracy (Supplementary Table 1) as the main accuracy assessment metrics. At the national level, we compared forest disturbance detected in this study to available existing dataset. Specifically, we compared the area of forest fire disturbance between our study and the national fire records during 2003–2009 (Supplementary Fig. 5). We compared the disturbance rates between our study and Landsat-derived global forest cover changes from 2001 to 20193 (Supplementary Fig. 4).Post-processingWe applied a series of spatial filters to minimize the unrealistic outliers from two potential sources of uncertainty, including speckle in time-series spectral trajectories or misregistration among images. This may lead to individual pixel or small patches including only a few pixels, which were (a) detected as disturbances, thus increasing the commission errors, or (b) not detected as disturbances, while their surrounding pixels were mostly disturbed, thereby increasing the omission errors. To address the issue (a), we removed all single-pixel disturbance patches through setting the minimum mapping unit as two 30 × 30 m2 pixels (0.18 ha). To address the issue (b), we applied a 3 by 3 moving window to fill holes through assigning the year of disturbance based on the years in the surrounding pixels. Finally, we smoothed the year of disturbance by assigning the center pixel using majority rules from surrounding pixels within the 3 by 3 windows, thus accounting for artefacts associated with uncertainties in the correct identification of the disturbance year.Characterizing disturbance regimes and their trendsWe characterized the disturbance regime using five indicators within each 0.5° grid cell (n = 1946) across China’s forests based on annual forest disturbance maps generated from the previous step. Within each grid cell, we calculated (1) total annually disturbed forest area (km2 yr−1), (2) percentage of forest disturbed annually (% yr−1), as annual disturbed forest area divided by the total forested area, (3) disturbance size (ha), as the number of disturbed pixels for each individual patch using an eight-neighbor rule, (4) disturbance frequency (# of patches per 1000 km2 forested area each year), as the number of disturbance patches per year divided by the total forested area, (5) disturbance severity (ΔNDVI = NDVIt−1 − NDVIt+1), as magnitude of NDVI change 1 year before and 1 year after disturbance, obtained from the LandTrendr analysis. We used (1) and (2) to characterize the disturbance rate, and (3)–(5) to describe the patch characteristics. The (2) and (4) were normalized by forest area within each grid cell, thus making them comparable among grid cells. For (3)–(5), we only calculated the patch size >0.45 ha (five 30 × 30-m2 pixels), because patches  TC2000), and the expansion of forested area from 1986 to 2000 (e.g., TC1986  20% following Liu et al., (2019). We should note that our study area did not include the newly afforested area after 2000. All analyses were performed within the forest mask, thus excluding the potential confounding factors from other land cover types. The description of TC1986 and TC2000 can be found in3,32. More

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Plant Ecology and Nature Conservation Group, Wageningen University, Wageningen, the NetherlandsFons van der Plas & Liesje MommerSystematic Botany and Functional Biodiversity, Life Science, Leipzig University, Leipzig, GermanyThomas Schröder-Georgi, Alexandra Weigelt, Kathryn Barry & Christian WirthGerman Centre for Integrative Biodiversity Research Halle-Jena-Leipzig, Leipzig, GermanyAlexandra Weigelt, Kathryn Barry, Adriana Alzate, Nico Eisenhauer, Anke Hildebrandt, Christiane Roscher & Christian WirthTerrestrial Ecology Research Group, School of Life Sciences Weihenstephan, Technical University of Munich, Munich, GermanySebastian Meyer & Wolfgang WeisserAquaculture and Fisheries Group, Wageningen University and Research Centre, Wageningen, the NetherlandsAdriana AlzateAgroécologie, AgroSup Dijon, Institut National de la Recherche Agronomique, Université de Bourgogne, Université de Bourgogne Franche-Comté, Dijon, FranceRomain L. BarnardEidgenössische Technische Hochschule Zürich, Zurich, SwitzerlandNina BuchmannDepartment of Experimental Plant Ecology, Institute for Water and Wetland Research, Radboud University Nijmegen, Nijmegen, the NetherlandsHans de KroonInstitute of Ecology and Evolution, University Jena, Jena, GermanyAnne Ebeling & Winfried VoigtInstitute of Biology, Leipzig University, Leipzig, GermanyNico EisenhauerHumboldt-Universität zu Berlin, Berlin, GermanyChristof EngelsInstitute of Plant Sciences, University of Bern, Bern, SwitzerlandMarkus FischerMax Planck Institute for Biogeochemistry, Jena, GermanyGerd Gleixner, Ernst-Detlef Schulze & Christian WirthHelmholtz Centre for Environmental Research, Leipzig, GermanyAnke HildebrandtFriedrich Schiller University Jena, Jena, GermanyAnke HildebrandtGeoecology, University of Tübingen, Tübingen, GermanyEva Koller-France & Yvonne OelmannInstitute of Geography and Geoecology, Karlsruhe Institute of Technology, Karlsruhe, GermanySophia Leimer & Wolfgang WilckeEcotron Européen de Montpellier, Centre National de la Recherche Scientifique, Montferrier-sur-Lez, FranceAlexandru MilcuCentre d’Ecologie Fonctionnelle et Evolutive, Unité Mixte de Recherche 5175 (Centre National de la Recherche Scientifique-Université de Montpellier-Université Paul-Valéry Montpellier-Ecole Pratique des Hautes Etudes), Montpellier, FranceAlexandru MilcuDepartment of Evolutionary Biology and Environmental Studies, University of Zurich, Zurich, SwitzerlandPascal A. NiklausUFZ, Helmholtz Centre for Environmental Research, Department Physiological Diversity, Leipzig, GermanyChristiane RoscherInstitute of Landscape Ecology, University of Münster, Münster, GermanyChristoph ScherberCentre for Biodiversity Monitoring, Zoological Research Museum Alexander Koenig, Bonn, GermanyChristoph ScherberGeobotany, Faculty of Biology, University of Freiburg, Freiburg, GermanyMichael Scherer-LorenzenCentre of Biodiversity and Sustainable Land Use, University of Göttingen, Göttingen, GermanyStefan ScheuJ.F. Blumenbach Institute of Zoology and Anthropology, Animal Ecology, University of Göttingen, Göttingen, GermanyStefan ScheuDepartment of Geography, University of Zurich, Zurich, SwitzerlandBernhard SchmidInstitute of Ecology, College of Urban and Environmental Sciences, Peking University, Beijing, ChinaBernhard SchmidLeuphana University Lüneburg, Institute of Ecology, Lüneburg, GermanyVicky TempertonAgroecology, Department of Crop Sciences, University of Göttingen, Göttingen, GermanyTeja TscharntkeF.v.d.P. wrote the initial draft of the manuscript. T.S.-G., A.W., K.B., S.M., A.A., R.L.B., N.B., H.d.K., A.E., N.E., C.E., M.F., G.G., A.H., E.K.-F., S.L., A.M., L.M., P.A.N., Y.O., C.R., C.S., M.S.-L., S.S., B.S., E.-D.S., V.T., T.T., W.V., W. Weisser, W. Wilcke and C.W. helped edit the manuscript. More

Here, we examine infection dynamics in the four regions of focus to learn more about how various control interventions performed in each region. Since the first documented cases emerged in these regions, the virus was able to spread freely across much of the first and second phases with a gradual increase in the control interventions. In the four regions of Makkah, Madinah, Eastern, and Riyadh, cases peaked on 12th May (6397 cases; 95% CI 5960–9697), 15th May (1967 cases; 95% CI 1625–2308), 23rd June (10367; 95% CI 8948–11785) and 11th June (11273 cases; 95% CI 11068–12491) respectively according to the fitted model shown in Fig. 2. As the epidemic progressed, more measures were adopted to contain the disease, and the disease’s infectiousness sharply decreased after the third period. There are a few factors responsible for the sudden declining trend: first our model is dependent on official data on cases that have been documented, and these data will only ever reflect a portion of the overall number of cases. Second, different regions developed different testing strategies, and some locations altered their approach to testing during the course of the time period that was investigated. It is possible that the beginning of the Hajj term (period 4) was a contributing factor in the decrease in the number of documented cases. Additionally, previous to this time period, the government indicated that it would be increasing the size of its local testing in order to detect new cases. It is possible that the efficacy of interventions would be reduced if increases are found to be occurring during the falling phase of an epidemic. This may result in measures being kept in place for a longer period of time than they would have been had more accurate data been provided.Figure 2With the use of the Delay Rejection Adaptive Metropolis method, the relevant parameters were estimated for each of the four areas of interest by fitting the data from 13th March until 25th September.Full size imageWe estimate the effective reproduction number (R_t) as an indicator of SARS-CoV-2 transmission before and after the interventions. Figure 3 depicts the dramatic shift in the rate of SARS-CoV-2 transmission as a result of decreased social contact and other control measures. At the beginning of the pandemic, (R_t) for SARS-CoV-2 in Saudi regions was between 4 and 6 as illustrated in Tables 6 and 7. In other words, on average each case spread to between four and six others. Considering that each new generation of SARS-CoV-2 cases occurs every five days, it is evident that this pandemic was rapidly expanding out of control. Moreover, we assumed that the transmission rate and the documented infection rate did not change during the first two periods since interventions were carried out gradually until a complete lockdown took place. As more measures were introduced, the spread of the disease began to decrease. Therefore, our data were based on the weekly reported number of documented SARS-CoV-2 cases broken down by region. As a result, it became clear that the reliability of the (R_t) value was relatively high for transmission.Figure 3Distribution of Rt estimates derived from 10000 MCMC samples for Makkah, Madinah, Eastern, and Riyadh, respectively. The black dot in the centre of each violin plot denotes the median, the thick bar in the plot denotes the interquartile range, and the thin bar in the plot denotes the lowest and maximum values. The mean and the credible interval for 95%, which is shown in parentheses, are labelled below or above, respectively.Full size imageThe effects of the events and interventions on the dynamics of SARS-CoV-2 in the regions of interest are considered. First, if the controls remained in phase four in Makkah, our model projects that the total number of documented cases would increase to 81047 (95% CI 79421–82672). In Al-Madinah, the cumulative number of documented cases would have increased to 22997 (95% CI 19578–26415). The number of cumulative documented cases may have reached 80520 (95% CI 78335–82704) if controls stayed steady in the Eastern region at the level they were at in phase four. If the pattern shown during the fourth period is taken into account, we estimate that there would have been 67150 (95% CI 63731–70568) documented infections in the Riyadh region. Figure 4 illustrates these findings.Figure 4The relevant parameters were estimated for each of the four regions of interest by first fitting the data of each region, and then predicting using the parameters from period 4. This was done with each of the four regions of interest separately.Full size imageWe now explore the impact of controls remaining in place at the same level as that implemented in phase three. In that case, the number of documented cases in Makkah would have increased to 116641 (95% CI 105015–128266). Similarly, the total number of documented cases in Al-Madinah would have increased to 53877 (95% CI 50458-57295) if the outbreak had been allowed to continue at the same level. If the controls had remained unchanged from how they were in phase three in the Eastern region, the total number of documented cases would have been 310459 (95% CI 298362–334981). Finally, in Riyadh this would have resulted in 665241 documented cases (95% CI 651822 to 678659). Figure 5 highlights these findings.Figure 5For each of the four areas of interest, the relevant parameters were estimated by first fitting the data of each region and then predicting using the parameters from period 3. This was carried out for each of the four areas of interest independently.Full size imageWe now investigate the impact of second-period controls remaining in place. In that case, the number of documented cases would increase to 1236642 (95% 1218314–1251626), 442865 (95% CI 439446–456283), 454031 (95% CI 441846–466215), and 2322624 (95% CI 1919206-3026042) in the regions of Makkah, Madinah, Eastern, and Riyadh, respectively (see Fig. 6).Figure 6For each of the four regions of interest, the relevant variables were identified by first fitting the data from each area and then making predictions using the parameters from period 2. Each of the four regions of interest was done separately.Full size imageThe efficacy of NPIs is dependent on when they are adopted, with earlier adoption resulting in greater success in lowering transmission rates of infectious diseases. In the early stages of COVID-19, Saudi regions made the decision to gradually implement measures in order to understand the severity of the disease and reduce the economic and social costs of lockdowns, as well as the political costs. In Fig. 6, if the government were to rely on the interventions of the second phase, then the number of cases of infection would considerably rise owing to the ineffectiveness of the measures. In the third period as in Fig. 5, the government made it possible to relax some of the control measures, but it is ultimately up to each area to decide whether they will maintain the same level of control or whether they will increase or decrease it. In comparison to the control measures carried out during the third and fourth periods, this led to significantly improved outcomes. The reason that these time periods were chosen is that there was no stiffening of the NPI response in most Saudi regions during the first two periods and control interventions were improved later on.Significant undetected infections resulted in the fast spread of new coronaviruses (SARS-CoV-2) which is illustrated in Fig. 7. The proportion of undocumented infections, including asymptomatic cases and undocumented symptomatic individuals who did not seek medical treatment or be tested for mild symptoms, was greater than that of Wuhan at the onset of the pandemic26, which may be a result of the following factors: first, the medical configuration was not optimal and public awareness was limited during the onset of the pandemic while the undocumented rate progressively increased; Second, contact tracing procedures employed in Saudi regions may have become overwhelmed if the number of early-stage cases in Saudi regions rises substantially. The discrepancy between the predicted proportions of asymptomatic (undocumented) cases may be attributable to the difficulty in the un-identifiability of parameters in epidemiological models. There were a substantial number of asymptomatic infected individuals with high infectivity in Saudi regions, where the epidemic situation escalated rapidly. Our research emphasises the frequency of asymptomatic SARS-CoV-2 cases and their role in transmission in order to increase people’s knowledge of asymptomatic cases and to serve as a guide for the prevention and control of SARS-CoV-2.Table 3 Estimated transmission rate in Saudi Regions.Full size tableTable 4 Estimated ascertainable infection rate.Full size tableIn this model, we fitted dynamic transmission rates because of varied preventable measures by the Saudi government at the level of the country or region. After a series of actions taken by the government, regions and cities went into lockdown, resulting in a decrease in the transmission rate as in Table 3. Before the interventions were introduced, in the first two periods of our study, we assumed the transmission rate did not change since individual and community responses had not effectively taken place. After severe interventions were implemented, the transmission rates were allowed to vary in later periods and reduced gradually due to the control measures that reduced the spread of disease27. Estimates of documented infection rates are presented in Table 4. Our model estimates show the documented infection rate has continued to decrease in the last two periods. Thus, the parameters we fit across periods are a measure of how effective the lockdown was in bringing down the documented infection rate28.Risk of resurgenceThe risk of resurgence in Saudi Arabia’s four regions has been examined in this section after the relaxation of intervention measures. There will be a rise in disease activity if control measures are relaxed without taking into account increases in the number of cases being detected, isolated, and/or traced. We predict the first week of no new cases of infection and the week when all current infections in Saudi Arabia will be eradicated.In the Makkah region, had the trend continued into the fourth period, the number of documented infections would have dropped to zero on average by the 6th September (23rd August to 27th September), and all infections would have been eradicated by the 26th of October (7th October to 14th November). On the 28th June, the number of weekly active infections (including presymptomatic, symptomatic, and asymptomatic cases) reached its highest point of 230,230 (95% CI 226811–234364), and on 8th September, that number dropped to 44023 (95% CI 40604–47441).Therefore, the number of documented infections would have reached zero in Al-Madinah region on average on 6th November (23rd October to 22nd November), and all infections would have been eliminated by 1st December (27th November to 14th December). On 23rd June, weekly active infections (including presymptomatic, symptomatic, and asymptomatic cases) peaked at 130,134 (95% CI 126715–133552) and then declined to 60023 (95% CI 58604–63441) on 25th September.If the trend had continued as it did in the fourth period in the Eastern region, the average number of documented infections would have reached zero on 2nd November (from 23rd October to 18th November), and the total eradication of infections would have happened on 1st December (26th November to 22nd December). The number of weekly active infections (including presymptomatic, symptomatic, and asymptomatic cases) peaked at 65000 (95% CI 61581–68418) during the week of July 23rd and subsequently decreased to 800 (95% CI 765–834) on 8th of September.Lastly, the model predicted that the number of weekly active infections in the Riyadh region (including presymptomatic, symptomatic, and asymptomatic infections) peaked on 28th June at 562332 (95% CI 513379–619542) and then decreased to 188215 (95% CI 174796–191633) on 18th September. On average, we expected that the number of documented infections would have decreased to zero on 18th October (7th October to 14th November) and that the total number of infections would have been eliminated on 1st December if the trend continued as it did in the fourth period (20th November to 23rd December). Figure 7 illustrates these findings. We found that if control measures were lifted 30 days following the first day of zero documented cases.Figure 7The estimated number of infected cases that were active (presymptomatic, symptomatic, and asymptomatic) during the research period in the areas of Makkah, Madinah, Eastern, and Riyadh respectively.Full size imageThe probability of resurgence, which we define as the number of active documented cases greater than 100 could be as high as 0.96 in Eastern, 0.95 in Madinah, 0.97 in Makkah, and 0.96 in Riyadh. If we adopt more stringent conditions of lifting controls after observing no confirmed cases for a continuous period of 30 days, the probability of resurgence decreases to 0.31, 0.28, 0.30, and 0.30, with probable resurgence occurring on 13th February, 7th February, 2nd January, and 8th January for Eastern, Makkah, Madinah and Riyadh, respectively (Fig. 8). Despite the use of a simplified model, these results emphasize the hazards of ignoring undetermined occurrences when modifying intervention techniques.Figure 8Figure demonstrating the effect of relaxing all control measures in all four regions 30 days following the first day without confirmed cases.Full size imageSensitivity analysisFor the purpose of testing the robustness of our research results, we conducted a series of sensitivity analyses by varying the durations of the latent and infectious periods, the ratio of transmissibility in asymptomatic (undocumented) cases to symptomatic (documented) cases, and the initial documented infection rate. We conduct eight sensitivity analyses (S1 to S8) within each model for each region of Saudi Arabia to assess the robustness of our model results. For instance, the sensitivity analysis performed for S1 was based on the changes of the latent period and pre-symptomatic infectious period, respectively, and other parameters remain the same. These modifications were carried out with the help of reference15,29, and the same approaches were used for the other parts of the sensitivity analysis, which is summarised in Table 5.Table 5 Description of essential model parameters that were not fitted in the MCMC, where (D_e) refers to the latent period, (D_p) refers to the pre-symptomatic infectious period, (D_i) refers to the symptomatic infectious period, (gamma _0) refers to the initial ascertain rate and (alpha) refers to the ratio of the transmission rate for P and A to I.Full size tableIn particular, for (S1), we raised the incubation period to 7 days (upper 95% CI based on ref15) and the pre-symptomatic infectious period to 3 days (upper 95% CI based on ref29). Therefore we set (D_e = 4) and (D_p=3), and modified (E_0) and (P_0) as needed. The transmissibility of the undocumented cases was assumed to be 0.46 (lower 95 % CI according to ref.31) of the infection cases for (S2); for (S3), the transmissibility of the asymptomatic (undocumented) cases was assumed to be 0.62 (upper 95 % CI according to ref31). We assumed that in (S4), the initial documented infection rate was (gamma _0) = 0.14 (lower 95 % CI according to ref13) and adjusted (A_0), (P_0) and E(0) accordingly. Similarly for (S5) we assumed the initial documented infection rate was (gamma _0) = 0.42 (upper 95 % CI according to ref13) and adjusted (P_0), (A_0), and (E_0) accordingly. In (S6) we set the variables (D_ e=3) and (D_p=1.1), and altered the values of (P_0) and (E_ 0) as necessary in accordance with13. In (S7) we assumed that the transmission rate of asymptomatic (undocumented) cases was half that of documented cases by setting 0.5. Finally, in (S8) we assumed that the infectious period ((D_i)) was double that of symptomatic cases by setting 6 days. Both (S7) and (S8) were based on30. The results of our sensitivity analysis are summarised in Tables 6 and 7. We note that the variation in the model predictions of (R_t) varies from setting to setting. However, these variations appear to be fairly small, proposing the robustness of the results to the specification of associated values in fairly realistic ranges13,13. Our sensitivity analysis provides information about the importance of each parameter to the model representing the transmission of SARS-CoV-2. An increase (or decrease) in parameter values, while other parameters’ values remain the same, contributes to an increase (or decrease) in effective reproduction numbers. For example, an increase in infectious period would result in a higher effective reproduction number at the beginning of the epidemic and a longer time required to clear all infections in Saudi regions32. Our sensitivity analysis indicates that almost all model parameters may have an important role in spreading this virus among susceptible people. In particular, the contact rate from person-to-person and the transition rate of asymptomatic (undetected) individuals play a significant role in disease spread. Our important findings, of a significant decrease in (R_t) after interventions and the existence of a substantial number of presymptomatic and asymptomatic cases, were found to be robust. This highlights that Saudi authorities should pay attention to intervention strategies in the event of a resurgence of cases and quarantining those who were in contact with active cases can effectively reduce the disease33. In Tables 6 and 7 we show the estimated effective reproduction number (R_t) associated with 95% CIs obtained from those eight sensitivity analyses for all four regions and all five time periods.Table 6 Sensitivity analysis of the effective reproduction number (R_t) for Eastern and Madinah.Full size tableTable 7 Sensitivity analysis of estimated effective reproduction number (R_t) for Makkah and Riyadh.Full size table More

We acknowledge members of the EDGE lab at the University of Gothenburg for stimulating discussions that inspired this comment. Special thanks to A. Bjorkman and R. Bjork for their feedback on the manuscript and to K. Happonen for help with data collection. More

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Evapotranspiration measurements and above-ground biomassFigure 1 shows daily evapotranspiration (ET, mm day−1) of each CC tested before mowing (DOY, day of the year, 184) and at 2, 8, 17 and 25 days after mowing (DOY 190, 196, 205 and 213); bare soil was also included as a reference. Before mowing, ET rates showed significant differences between and within the three groups. CR plants had a mean ET of 8.1 mm day−1, which was lower, compared to the other two groups (10.6 and 18.6 mm day−1 for GR and LE, respectively) and the bare soil control (8.5 mm day−1). On DOY 184, values as high as 9.4 (Glechoma hederacea L., GH) and 9.8 mm day−1 (Trifolium subterraneum L. cv. Denmark, TS) were found (Fig. 1), while ranging around 7 mm day-1, Dichondra repens J.R.Forst. & G.Forst. (DR), Hieracium pilosella L. (HP), and Sagina subulata (Swartz) C. Presl (SS) ET were lower than soil evaporation itself.Figure 1Vertical bars represent the daily water use as referred to unit of soil (ET, mm day−1) for the bare soil (yellow) and all the cover crop species as divided into creeping plants (shades of blue), legumes (shades of green) and grasses (shades of orange). Evapotranspiration was measured though a gravimetric method before (i.e. − 4) and at 2, 8, 17 and 25 days after mowing. ET data are mean values ± SE (n = 4).Full size imageOn the same day, a large ET variation was recorded within the GR group as Festuca arundinacea Schreb. cv. Thor (FA) scored the highest daily ET values (13.4 mm day−1), whereas in Festuca ovina L. cv. Ridu (FO), water loss was reduced by 45% (7.5 mm day−1). Within the 15 CCs, LE registered the highest pre-mowing ET with Trifolium michelianum Savi cv. Bolta (TM) peaking at 22.6 mm day−1. However, within LE, Medicago polymorpha L. cv. Scimitar (MP) showed ET values as low as 12.1 mm day−1 (Fig. 1).Two days after mowing, all tested CCs recorded ET values lower than 9 mm day−1 (Fig. 1). Moreover, water use reduction among LE ranged between 56% (M. polymorpha, MP) and 73% (T. michelianum, TM), such that T. michelianum (TM, 6.1 mm day−1), Medicago truncatula Gaertn. cv. Paraggio (MT, 5.6 mm day−1) and M. polymorpha (MP, 5.2 mm day−1) registered ET values lower than the bare soil (7.0 mm day−1). Even though registering a consistent ET reduction after mowing, GR retained ET rates slightly higher than bare soil, except for F. ovina (FO), which recorded the lowest at 6.3 mm day−1. Subsequent samplings showed that most of the CCs had a progressive recovery in water use (Fig. 1) and data taken 17 days after mowing confirmed that Lotus corniculatus L. cv. Leo (LC) and all GR fetched pre-mowing ET rates. Medicago lupulina L. cv. Virgo (ML) registered a partial recovery with similar rates (about 13 mm day−1) at 17 and 25 days after the mowing event. F. ovina and all remaining LE stayed below 10 mm day−1 with ET values close to the control until the end of the trial. At 17 days from grass cutting, under a quite high exceeding-the-pot biomass, both G. hederacea (GH) and T. subterraneum (TS) reached ET values as high as 12.0 and 11.4 mm day−1, respectively. On the other hand, D. repens (DR), H. pilosella (HP), and S. subulata (SS) even though with slightly higher ET values than those registered at the beginning of the trial (DOY 184), remained close to the soil evaporation rates until DOY 213.Aboveground dry clipped biomass at the first mowing date (ADW_MW1, DOY 188) showed large differences among groups, as represented in Table 1. ADW_MW1 within LE was quite variable, as values ranged between 274.3 g m−2 (M. polymorpha, MP) and 750.0 g m−2 (T. michelianum, TM). With a mean value of 565.9 g m−2, LE aboveground biomass was 80% higher than the mean GR ADW_MW1 (110.2 g m-2). F. ovina (FO) scored the lowest value at 48.4 g m−2 among grasses, while within the creeping group, G. hederacea (GH) and T. subterraneum (TS) had biomass development outside the pot edges totalling 89.6 g m−2 and 23.2 g m−2, respectively.Table 1 Aboveground dry biomass clipped at the first mowing event (ADW _MW1), the corresponding leaf area surface index (LAI) and water use per leaf area unit (ETLEAF) of all cover crops tested.Full size tableLeaf area index (LAI, m2 m−2) at mowing showed the highest values in LE with LAI peaking at 12.4 (Table 1). Among GR, LAI did not show significant differences, being around 1.2. Concerning CR, LAI was assessed at 0.2 and 0.8 for T. subterraneum (TS) and G. hederacea (GH) respectively, while LAI estimated through photo analysis ranged between 1.3 (D. repens, DR) and 3.6 (T. subterraneum TS).Evapotranspiration per leaf area unit (ETLEAF) was notably higher in GR, ranging between 7.75 (F. ovina, FO) and 9.22 (Lolium perenne L. cv. Playfast, LP) mm m−2 day−1 (Table 1). In descending order, ETLEAF was the highest in D. repens (DR, 5.46 mm m−2 day−1). Similar ETLEAF was found when comparing some LE and CR species such as M. truncatula (MT, 3.40 mm m−2 day−1), M. lupulina (ML, 4.05 mm m−2 day−1), G. hederacea (GH, 3.68 mm m−2 day−1), H. pilosella (HP, 3.86 mm m-2 day-1) and T. subterraneum (TS, 2.74 mm m−2 day−1). T. michelianum (TM), with 1.81 mm m-2 day-1 scored the lowest ETLEAF of all species (Table 1).Plotting LAI versus the before-mowing ET yielded a significant quadratic relationship (R2  > 0.76) (Fig. 2a) which helped to distinguish two different data clouds. Till LAI values of about 6, the model was linear, having at its lower end all GR and CR species with the inclusion of M. polymorpha (MP) as a legume, while, at the other end, M. truncatula (MT), L. corniculatus (LC) and M. lupulina (ML) were grouped together. T. michelianum (TM) was isolated from all CCs at 22.56 mm day−1.Figure 2Panel (a): quadratic regression of leaf area index (LAI, m2 m−2) vs cover crop evapotranspiration per unit of soil (ET, mm day−1). Each data point is mean value ± SE (n = 4). The quadratic model equation is y = − 0.128×2 + 2.9968x + 5.4716, R2 = 0.76. Panel (b): the quadratic regression between LAI corresponding to the clipped biomass (m2 m−2) and cover crop ET reduction (%). Each data point is mean value ± SE (n = 4). Quadratic model equation is y = − 0.8985×2 + 16.503x + 5.1491, R2 = 0.94.Full size imageWhen regressing the fraction of ET reduction, compared to pre-mowing values vs LAI (Fig. 2b), the same quadratic model achieved a very close fit (R2 = 0.94, p  1 mm) root diameters as affected by soil cover.Full size tableThe highest values of diameter class length (DCL, mm cm−3) for very fine roots (DCL_VF,  1.0 mm) roots although, most notably, L. corniculatus roots showed the highest abundance for both DCL_M (23.08 cm cm−3) and DCL_C (0.54 cm cm−3).At the 10–20 cm soil depth, GR confirmed the highest values for both very fine and fine roots, with F. arundinacea reaching maximum DCL of 2.269 and 5.215 cm cm-3, respectively (Table 2). L. corniculatus largely outscored any other species for both medium and coarse root diameter (6.173 and 0.037 cm cm−3, respectively), with F. arundinacea ranking second (3.157 and 0.016 cm cm−3, respectively).The highest root dry weight (RDW, mg cm-3) within the topsoil layer was reached by L. corniculatus (8.7 mg cm−3) and F. arundinacea (7.6 mg cm-3). Notably, such values were significantly higher than those recorded on the remaining species, except for the F. arundinacea vs F. rubra commutata comparison (Table 2). At 10–20 depth, scant variation was recorded in RDW measured in grasses, whereas L. corniculatus held its supremacy within legumes (4.5 mg cm−3). Within the creeping type, D. repens (DR) and G. hederacea (GH) scored RDW values as high as those determined for grass species (namely F. arundinacea , P. pratensis and F. rubra commutata), whereas S. subulata (SS) essentially had no root development.Soil aggregates and mean weight diameter (MWD)Table 3 reports the proportional aggregate weight (g kg−1) for both 0–10 and 10–20 cm soil depths. Compared to bare soil, the largest increase in large macroaggregates (LM,  > 2000 µm) in the top 10 cm of soil was achieved by L. corniculatus with 461 g kg−1. L. corniculatus differed from the rest of the LE group, whose grand mean (90 g kg−1) was the lowest of the three tested groups. As a legume, T. subterraneum (TS, 122 g kg−1) recorded the lowest values compared to fellow CR species, ranging between 211 (D. repens, DR) and 316 g kg−1 (G. hederacea, GH). GR recorded LM values slightly lower than those of CR, with a mean value of 217 vs 224 g kg-1.Table 3 Proportional aggregate weight (g kg−1) of sand-free aggregate-size fractions acquired from wet sieving as affected by soil cover and mean weight diameter (MWD). Aggregate-size fraction divided as macroaggregates with large size ( > 2 mm, LM) and small size (2 mm—250 μm, sM), microaggregates (250 μm—53 μm, m), and silt and clay ( More