Philippa Kaur

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Literature search and study selectionA systematic literature search was conducted in the ISI Web of Science database for observational and experimental studies published from 1975 to 13 January 2020 using the following search terms: TOPIC: (grassland* OR prairie* OR steppe* OR shrubland* OR scrubland* OR bushland*) AND TOPIC: (drought* OR ‘dry period*’ OR ‘dry condition*’ OR ‘dry year*’ OR ‘dry spell*’) AND TOPIC: (product* OR biomass OR cover OR abundance* OR phytomass). The search was refined to include the subject categories Ecology, Environmental Sciences, Plant Sciences, Biodiversity Conservation, Multidisciplinary Sciences and Biology, and the document types Article, Review and Letter. This yielded a total of 2,187 peer-reviewed papers (Supplementary Fig. 1). At first, these papers were screened by title and abstract, which resulted in 197 potentially relevant full-text articles. We then examined the full text of these papers for eligibility and selected 87 studies (43 experimental, 43 observational and 1 that included both types) on the basis of the following criteria:

(1)

The research was conducted in the field, in natural or semi-natural grasslands or shrublands (for example, artificially constructed (seeded or planted) plant communities or studies using monolith transplants were excluded). We used this restriction because most reports on observational droughts are from intact ecosystems, and experiments in disturbed sites or using artificial communities would thus not be comparable to observational drought studies.

(2)

In the case of observational studies, the drought year or a multi-year drought was clearly specified by the authors (that is, we did not arbitrarily extract dry years from a long-term dataset). Please note that some observational data points are from control plots of experiments (of any kind), where the authors reported that a drought had occurred during the study period. We did not involve gradient studies that compare sites of different climates, which are sometimes referred to as ‘observational studies’.

(3)

The paper reported the amount or proportion of change in annual or growing-season precipitation (GSP) compared with control conditions. We consistently use the term ‘control’ for normal precipitation (non-drought) year or years in observational studies and for ambient precipitation (no treatment) in experimental studies hereafter. Similarly, we use the term ‘drought’ for both drought year or years in observational studies and drought treatment in experimental studies. In the case of multi-factor experiments, where precipitation reduction was combined with any other treatment (for example, warming), data from the plots receiving drought only and data from the control plots were used.

(4)

The paper contained raw data on plant production under both control and drought conditions, expressed in any of the following variables: ANPP, aboveground plant biomass (in grassland studies only) or percentage plant cover. In 79% of the studies that used ANPP as a production variable, ANPP was estimated by harvesting peak or end-of-season AGB. We therefore did not distinguish between ANPP and AGB, which are referred to as ‘biomass’ hereafter. We included the papers that reported the production of the whole plant community, or at least that of the dominant species or functional groups approximating the abundance of the whole community.

(5)

When multiple papers were published on the same experiment or natural drought event at the same study site, the most long-term study including the largest number of drought years was chosen.

In addition to the systematic literature search, we included 27 studies (9 experimental, 17 observational and 1 that included both types) meeting the above criteria from the cited references of the Web of Science records selected for our meta-analyses, and from previous meta-analyses and reviews on the topic. In total, this resulted in 114 studies (52 experimental, 60 observational and 2 that included both types; Supplementary Note 9, Supplementary Fig. 2 and ref. 25).Data compilationData were extracted from the text or tables, or were read from the figures using Web Plot Digitizer26. For each study, we collected the study site, latitude, longitude, mean annual temperature (MAT) and precipitation (MAP), study type (experimental or observational), and drought length (the number of consecutive drought years). When MAT or MAP was not documented in the paper, it was extracted from another published study conducted at the same study site (identified by site names and geographic coordinates) or from an online climate database cited in the respective paper. We also collected vegetation type—that is, grassland when it was dominated by grasses, or shrubland when the dominant species included one or more shrub species (involving communities co-dominated by grasses and shrubs). Data from the same study (that is, paper) but from different geographic locations or environmental conditions (for example, soil types, land uses or multiple levels of experimental drought) were collected as distinct data points (but see ‘Statistical analysis’ for how these points were handled). As a result, the 114 published papers provided 239 data points (112 experimental and 127 observational)25.For the observational studies, normal precipitation year or years specified by the authors was used as the control. If it was not specified in the paper, the year immediately preceding the drought year(s) was chosen as the control. When no data from the pre-drought year were available, the year immediately following the drought year(s) (14 data points) or a multi-year period given in the paper (22 data points) was used as the control. For the experimental studies, we also collected treatment size (that is, rainout shelter area or, if it was not reported in the paper, the experimental plot size).For the calculation of drought severity, we used yearly precipitation (YP), which was reported in a much higher number of studies than GSP. We extracted YP for both control (YPcontrol) and drought (YPdrought). For the observational studies, when a multi-year period was used as the control or the natural drought lasted for more than one year, precipitation values were averaged across the control or drought years, respectively. Consistently, in the case of multi-year drought experiments, YPcontrol and YPdrought were averaged across the treatment years. When only GSP was published in the paper (63 of 239 data points), we used this to obtain YP data as follows: we regarded MAP as YPcontrol, and YPdrought was calculated as YPdrought = MAP − (GSPcontrol − GSPdrought). From YPcontrol and YPdrought data, we calculated drought severity as follows: (YPdrought − YPcontrol)/YPcontrol × 100.For production, we compiled the mean, replication (N) and, if the study reported it, a variance estimate (s.d., s.e.m. or 95% CI) for both control and drought. In the case of multi-year droughts, data only from the last drought year were extracted, except in five studies (17 data points) where production data were given as an average for the drought years. When both biomass and cover data were presented in the paper, we chose biomass. For each study, we consistently considered replication as the number of the smallest independent study unit. When only the range of replications was reported in a study, we chose the smallest number.To quantify climatic aridity for each study site, we used an aridity index (AI), calculated as the ratio of MAP and mean annual PET (AI = MAP/PET). This is a frequently used index in recent climate change research27,28. AI values were extracted from the Global Aridity Index and Potential Evapotranspiration (ET0) Climate Database v.2 for the period of 1970–2000 (aggregated on annual basis)29.Because we wanted to prevent our analysis from being distorted by a strongly unequal distribution of studies between the two study types regarding some potentially important explanatory variables, we left out studies from our focal meta-analysis in three steps. First, we left out studies that were conducted at wet sites—that is, where site AI exceeded 1. The value of 1 was chosen for two reasons: above this value, the distribution of studies between the two study types was extremely uneven (22 experimental versus 2 observational data points with AI  > 1)25, and the AI value of 1 is a bioclimatically meaningful threshold, where MAP equals PET. Second, we left out shrublands, because we had only 14 shrubland studies (out of 105 studies with AI  More

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Characteristics of heavy metal concentrationsOn the basis of the soil sample collection and chemical analysis, the concentration data for heavy metals in the urban soils of Wuxi were obtained. Through the statistical analysis of the soil heavy metal concentration data (Table 1), on the whole, the concentration of each heavy metal is as follows: Mn  > Zn  > Cr  > Ni  > Pb  > Cu  > Co  > Be  > Cd. Among these, the concentration range of Cr was 64.5–99 mg kg-1, and the average concentration was 72.9 mg kg−1. The concentration range of Ni was 31.4–67.5 mg kg−1, and the average concentration was 38.2 mg kg−1. The concentration range of Cu was 19.8–37.2 mg kg−1, and the average concentration was 25.5 mg kg−1. The concentration range of Zn was 72.4–1146 mg kg−1, and the average concentration was 90.2 mg kg−1. The concentration range of Cd was 0.34–1.06 mg kg−1, and the average concentration was 0.51 mg kg−1. The concentration range of Pb was 25.6–66.4 mg kg−1, and the average concentration was 37.6 mg kg−1. The variation coefficients of urban soil heavy metal concentration in Wuxi is between 0.09 and 0.33, which is less than 1. The spatial fluctuation of urban soil heavy metal concentration in Wuxi is small, indicating that the sources may be the same or similar.Table 1 Statistics of the heavy metal concentrations and Pb isotope ratios in the urban soils of Wuxi city (unit of heavy metal: mg kg−1; CV: coefficient of variation).Full size tableBy analysing the spatial distributions of the urban soil heavy metal concentrations in Wuxi, several obvious spatial distribution characteristics are found (Fig. 2). First, the heavy metals have high values in the central area of Wuxi, due to where has a high population density and various industries. The central aggregation of Pb is more obvious. Due to the dense roads in the city centre, vehicle traffic, bus stop signs and gas stations are mostly concentrated here, which will lead to Pb contents in this area that are significantly higher than those in other areas. In addition to the heavy metal concentrations, such as those for Cu, Zn and Cr in the downtown area, there are also areas with high values in western Wuxi and low values in eastern Wuxi. This phenomenon may be related to the land use types in Wuxi. In the western area of Wuxi, most land use types are urban and construction land, and the soils in this area are greatly disturbed by human activities. In the eastern region of Wuxi, woodland and grassland account for a large proportion of the land use types, which are less disturbed by human activities.Figure 2Spatial distribution characteristics of heavy metals in the urban soils of Wuxi city (unit: mg kg−1) [the figure was generated by Yan Li using the ArcGIS 10.2 (http:// https://developers.arcgis.com/)].Full size imageSource analysis of heavy metalsExploring for heavy metal pollution from emission sources is an important prerequisite for the study of urban soil pollution. By analysing the sources of heavy metals in soil environments, we can accurately determine which industries are major sources28,29,30 and whether there is homologous pollution. This is not only a theoretical basis for the study of lake sediment pollution and to clarify the risks brought by different pollution sources to the urban soil environment but also provides important guides for local government control of specific polluting industries and pollutant emissions. Based on this, the correlations and significance of heavy metals in the urban soils of Wuxi were analysed (Table 2). Generally, a heavy metal pollution source will emit multiple heavy metals at the same time. If the pollution source has a large emission, the concentration of these heavy metals in the environment will show a high level; on the contrary, if the emission of this pollution source is small, the concentration of these heavy metals in the environment will show a low level10. The correlations between the heavy metals Zn, Cr, Ni, Pb, Cu and Cd are between 0.655–0.907 and show strong correlations and significance at a level of 0.01. The strong significant correlations between different heavy metals indicate that these heavy metals have similar emission sources and transmission routes, which also means that they have consistent sources.Table 2 Correlations of Heavy Metals in the Urban Soils of Wuxi City.Full size tableTo further determine which industries are the sources of the heavy metals found in the urban soil of Wuxi, we analysed the Pb isotope data. The variation range of 208Pb/206Pb in soil is 2.09–2.12, and the average value is 2.10. The variation range of 206Pb/207Pb in soil is 1.17–1.18, and the average value is 1.177 (Table 1). After consulting relevant literature and materials, the main pollution sources of heavy metals in cities in eastern China include coal combustion, oil combustion, factory emissions, municipal wastes and so on3. Therefore, we collected the corresponding Pb isotope data in the emissions of heavy metal pollution sources. By collecting and comparatively analysing the Pb isotope data of known pollution sources (Fig. 3), it was determined that the Pb isotopes of the urban soil heavy metals in the soils of Wuxi have distinct characteristics. First, the Pb isotope distributions in the soils of Wuxi are relatively concentrated, and the ranges of variation are relatively small, which indicate that these heavy metals may have the same source or similar sets of sources. Second, the Pb isotopes in the urban soils of Wuxi city have few similarities with those of the uncontaminated soils and granites in eastern China; in contrast, the Pb isotopes in the urban soils of Wuxi are distributed in areas that are associated with coal combustion, automobile exhaust and urban waste (supplementary materials). The urban soil heavy metals in Wuxi generally have similar pollution sources and are greatly affected by human activities such as coal combustion and automobile exhaust emissions. Wuxi has a developed industrial economy and large numbers of factories. In the production and processing activities, the combustion of energy and fuel and the incomplete utilization of raw materials will lead to the enrichment of pollutants in the surrounding environment. By comparing other studies30,31, the Pb isotope analysis results in this study well indicate the source of soil heavy metals in Wuxi and make up for the Pb isotope data in this area. In the process of urban development, we should develop and apply clean energy, reduce the utilization of petroleum fossil fuels, and control the enrichment of heavy metals and other pollutants in the soil from the source.Figure 3Comparison of the Pb isotope compositions in the urban soils of Wuxi city with known sources.Full size imageEcological risk analysisBy calculating the potential ecological risk index for the heavy metals in the urban soils of Wuxi, the risks of heavy metals in the Wuxi soils were evaluated (Table 3). According to previous studies21, an Ei value lower than 40 indicates that a heavy metal is in a low-risk state at this location, and Ei values greater than or equal to 40 indicate that a heavy metal represents a high-risk state at this location. The average value of the potential ecological risk index of soil heavy metal Cd in Wuxi is 80.3, which represents a high-risk state. The average distributions of the potential ecological risk indexes of the heavy metals Cr, Cu, Zn, Pb and Ni are 1.8, 4.3, 1.1, 5.5 and 4.8, respectively, which all indicate a low-risk state. The risk statuses of different heavy metals may show certain correlations in space, which may be mutually complementary or antagonistic. Examining the spatial interactions of different heavy metal compound pollutants in urban soils plays an important role in the prevention and control of urban heavy metal pollution. Based on this, we used the Lisa analysis method to explore the spatial correlations of the different heavy metal risks in the urban soils of Wuxi (Fig. 4). The Moran scatter diagram can be divided into four quadrants that correspond to four different spatial patterns. High means that the variable value is higher than the average value, and Low means that the variable value is lower than the average value. In the upper right quadrant (High–High), a high-value area is surrounded by high-value neighbours; in the upper left quadrant (Low–High), a low-value area is surrounded by high-value neighbours; in the lower left quadrant (LL), a low-value area is surrounded by low-value neighbours; and in the lower right quadrant (High–Low), a high-value area is surrounded by low-value neighbours. High-High and Low-Low indicate that the differences between the region and its surrounding areas are small; that is, the regions with higher or lower values are concentrated, while the Low–High and High–Low quadrants indicate that the variable values between a region and its surrounding areas are different to a certain extent.Table 3 Ecological risk and health risk analysis of heavy metals in the urban soils of Wuxi (Cr-E represents the ecological risk of metal element Cr; Ni-E represents the ecological risk of metal element Ni; Cu-E represents the ecological risk of metal element Cu; Zn-E represents the ecological risk of metal element Zn; Cd-E represents the ecological risk of metal element Cd; Pb-E represents the ecological risk of metal element Pb; ADDderm-C is the average exposure to skin contact pathways for child; ADDderm-A is the average exposure to skin contact pathways for adult; ADDing-C is the average daily exposure to intake pathway for child; ADDing-A is the average daily exposure to intake pathway for adult; HI-C is the total health risk caused by accumulation of heavy metals in multiple ways in the same environmental medium for child; HI-A is the total health risk caused by accumulation of heavy metals in multiple ways in the same environmental medium for adult).Full size tableFigure 4LISA analysis of the ecological risks from different heavy metals [the figure was generated by Yan Li using the ArcGIS 10.2 (http:// https://developers.arcgis.com/)].Full size imageIn this study, two main results were obtained from spatial correlation Lisa analysis between different heavy metals. One is a High-High area, which is mainly distributed in the central and western regions of Wuxi city, which is consistent with the spatial distribution of the urban soil heavy metal concentrations in Wuxi city and is strongly disturbed by human activities. The other is the insignificant area, in which there are also large numbers of factories and enterprises and in which the forestland and grassland are distributed at intervals, which leads to an insignificant spatial correlation of soil heavy metal contents. Based on the above analysis, the high-risk areas for heavy metals in the urban soils of Wuxi are mainly concentrated in the central and western regions, and the relevant management activities need to be given great attention. In the eastern region, sporadic high-risk areas are also present, which should also receive due attention. Moran’s I is a method to measure the interdependence and degree of objects or phenomena by constructing statistics on certain characteristics or attributes for a certain spatial unit in the study area and the surrounding spatial units. It can be used to describe the spatial characteristics of spatial units such as aggregation or outliers in the distribution of certain attributes and is a very important technology in spatial data analysis33,34. However, few studies have applied it to the spatial relationship analysis of different heavy metals in urban soil.Health risk analysisBy using the health risk assessment model that is recommended by the U.S. EPA, this study calculated the health risks of soil heavy metals to adults and children through skin contact and ingestion. For both adults and children, the risk of soil heavy metals through ingestion was much higher than that caused by skin exposure (Table 3). For children, the total health risk that was caused by soil heavy metals is 0.078, which is four times that of adults. This may be related to children’s habits. Most children like to play with sand and climb around on the ground. These behaviours greatly increase the frequency of children contacting the soil, which thus increases the health risk caused by heavy metals in the soil. To further explore the spatial characteristics of the health risks of heavy metals in the soils of Wuxi, this study provides spatial predictions of the health risk values of soil heavy metals (Fig. 5). The total health risk values of soil heavy metals for children and adults have similar spatial distribution characteristics. High health risk values appear in the central area of Wuxi and decrease in a ring-shaped pattern. This is similar to the development degree of the city. The downtown area of Wuxi is densely populated, the pedestrian flow is very large, and the health risk of soil heavy metals in this area is very high, which poses a very serious potential threat. The health risk values for the western region of Wuxi are high, and there is also a potential threat. When compared with western Wuxi, eastern Wuxi has a lower risk.Figure 5Health risk analysis of heavy metals in the urban soils of Wuxi [the figure was generated by Yan Li using the ArcGIS 10.2 (http:// https://developers.arcgis.com/)].Full size image More

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