Abstract
The frequency and intensity of droughts have increased over the decades, leading to increased forest decline. The response of forest to drought can be evaluated by both its sensitivity to drought (resistance) and its post-drought recovery rate (resilience). However, it remains uncertain how drought resistance and resilience of forests change over time under climate change. We assessed the spatiotemporal dynamics of forest resistance and resilience to drought over the past century (1901â2015) with global tree ring data records from 2,935âsites, in conjunction with plant trait data. We found that gymnosperms and angiosperms showed different spatial patterns of drought resistance and resilience, driven by variations in eco-physiological traits. Resistance and resilience also varied with drought seasonal timing. Surprisingly, we found that the trade-off between resistance and resilience for gymnosperms, previously reported only spatially, also occurred at the temporal scale. In particular, drought resilience markedly increased, but resistance decreased, for gymnosperms between 1950â1969 and 1990â2009, indicating that previous model simulations assuming invariant resistance may have underestimated the impacts of drought on gymnosperm-dominated forests under future climate change.
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Data availability
All data from public sources used in this study can be obtained from https://figshare.com/articles/Trade-off_between_gymnosperm_resistance_and_resilience_increases_forest_sensitivity_to_extreme_drought/12047241.
Code availability
Computer codes for the analysis of the data are available from https://github.com/LilyXiangyi/ForestResponse.
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Acknowledgements
This study was supported by the National Natural Science Foundation of China (nos. 41530528 and 41988101) and the Second Tibetan Plateau Scientific Expedition and Research Program (no. 2019QZKK0208). P.C. and J.P. acknowledge support from European Research Council Synergy project SyG-2013-610028 IMBALANCE-P and P.C. acknowledges support from the ANR CLAND Convergence Institute. We gratefully acknowledge all voluntary researchers who have contributed to and maintained the ITRDB. We also thank A. K. Post for proofreading the article.
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S.P. designed the research. X.L. performed analysis and created all the figures. X.L. and S.P. drafted the paper. All authors contributed to interpretation of the results and to the text.
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Extended data
Extended Data Fig. 1 Examples for identifying season categories.
(a) Location of the ring-width index (RWI) at four sites, co551 (40.05°N, 105.43°W), chin048 (30.30°N, 91.52°E), zimb001 (18.67°S, 26.68°E), and ausl004 (32.00°S, 115.65°E). The variation of monthly precipitation minus potential evapotranspiration (P-PET) anomalies divided each drought year into a dry season and a wet season (see details of identification of season categories in Methods) for RWI sites (b) co551 in North America with an elevation of 2560m, (c) chin048 in Asia (elevation missing), (d) zimb001 in Africa with an elevation of 1085m, and (e) ausl004 in Australia with an elevation of 328m. The shaded areas in (b-e) indicate the dry season. The names of the four RWI sites are taken from the ITRDB tree-ring database.
Extended Data Fig. 2 Examples for identifying drought classification for a drought year using RWI site co551.
The drought threshold in (a) the dry season (ThresholdDS) and (b) the wet season (ThresholdWS), derived from the tenth percentile of the data distribution (see details of drought classification in Methods). c, 1966, a DS drought year with a drought only in the dry season, indicated by a mean SPEI in the dry season lower than the drought threshold in the dry season (SPEIDS < ThresholdDS) but a mean SPEI in the wet season higher than the drought threshold in the wet season (SPEIWS > ThresholdWS). d, 1981, a WS drought year with a drought only in the wet season, indicated by SPEIDS > ThresholdDS but SPEIWS < ThresholdWS. e, 1977, a DS+WS drought year with a drought in both the dry and wet seasons, indicated by SPEIDS < ThresholdDS and SPEIWS < ThresholdWS.
Extended Data Fig. 3 Mean resistance and resilience of gymnosperms and angiosperms in drought years.
a, resistance to drought in the dry season (DS droughts), (b) resilience to drought in the dry season (DS droughts), (c) resistance to drought in the wet season (WS droughts), (d) resilience to drought in the wet season (WS droughts). Circles represent the gymnosperm tree-ring sites, and triangles represent the angiosperm tree-ring sites. The color bars denote the range of magnitude of (a, c) mean resistance (Rt) and (b, d) mean resilience (Rs), respectively.
Extended Data Fig. 4 Comparison of ln-transformed resistance (ln(Rt)) and resilience (ln(Rs)) for drought years in both the dry and wet seasons (DS+WS drought), the dry season (DS drought), and the wet season (WS drought) for gymnosperms and angiosperms.
For each row, different letters within a row represent significant differences (p < 0.05) among the drought timings. Values shown in the table are all unitless.
Extended Data Fig. 5 Drought severity (indicated by SPEI) of WS, DS and DS+WS droughts at gymnosperm and angiosperm tree-ring sites.
Significant differences (p < 0.05) among the drought timings were identified by an analysis of variance (one-way ANOVA) and are marked by different letters for gymnosperms (denoted by filled boxplots) and for angiosperms (denoted by open boxplots). The boxes indicate the 25th and 75th percentiles, and the lines in the boxes indicate the medians.
Extended Data Fig. 6 Temporal changes of resistance and resilience for gymnosperm and angiosperm forests during 1910-2009.
Temporal change of (a) gymnosperm resistance, (b) angiosperm resistance, (c) gymnosperm resilience, (d) angiosperm resilience. 290 gymnosperm sites in (a) and (c) and 65 angiosperm sites in (b) and (d) experiencing droughts in all four periods, i.e. before 1950, 1950-1969, 1970-1989, and 1990-2009 were used for temporal analysis. The x-axis represents the range of ln-transformed resistance (ln(Rtâ+â1)) and resilience (ln(Rsâ+â1)) at each RWI site during three periods. Histograms (left y-axis) show the distribution of ln (Rtâ+â1) and ln(Rsâ+â1) and lines (right y-axis) show the kernel density distribution of ln(Rtâ+â1) and ln(Rsâ+â1) during the period of before 1950 (gray), 1950-1969 (blue), 1970-1989 (orange), and 1990-2009 (purple). The boxplots in the inset of each panel show the distribution of temporal change in 1970-1989 (orange) and 1990-2009 (purple) relative to 1950-1969 for all sites. The yellow lines represent median values and points represent mean values in the boxes. The percentages of sites showing positive change (+) and negative change (-) are displayed in the panels. Significant differences (p < 0.05) of all sites between 1970-1989 and 1950-1969, and between 1990-2009 and 1950-1969 were identified by Kruskal-Wallis test and denoted by asterisks (*).
Extended Data Fig. 7 Comparison of hydraulic safety margin (HSM) and specific leaf area (SLA) between gymnosperms and angiosperms.
a, Empirical cumulative distribution function (CDF) of HSM between gymnosperms (black) and angiosperms (blue). b, Empirical CDF of SLA between gymnosperms and angiosperms.
Extended Data Fig. 8 Spatial correlations of differences in resistance and resilience with differences in tree growth, temperature, precipitation, and drought severity between two consecutive periods.
a, Correlations between resistance and RWI (upper panel), and between resilience and RWI (lower panel) across gymnosperm (blue bars) and angiosperm (orange bars) forest sites. b, Correlations between resistance and drought severity (upper panel) and between resilience and drought severity (lower panel) across gymnosperm and angiosperm forest sites. c, Correlations between resistance and temperature (upper panel) and between resilience and temperature (lower panel) across gymnosperm and angiosperm forest sites. d, Correlations between resistance and precipitation (upper panel) and between resilience and precipitation (lower panel) across gymnosperm and angiosperm forest sites. I, differences between 1950-1969 and 1970-1989; II, differences between 1970-1989 and 1990-2009. Significance correlations are denoted by *** (p < 0.001), ** (p < 0.01), and * (p < 0.05).
Extended Data Fig. 9 An example of decrease in drought resistance with increase in drought severity.
A tree-ring site (32.43°N, 110.79°W, Ponderosa pine) in Arizona, USA presents a decrease in drought resistance with increase in drought severity during three periods (i.e. 1950-1969, 1970-1989, and 1990-2009). The name of the tree-ring site is az598 in the ITRDB tree-ring database. Blue bars (upper panel) denote the mean drought severity during each period and orange bars (lower panel) denote the mean resistance during each period. Arrows show the directions of changes in drought severity (blue) and resistance (orange) over time.
Extended Data Fig. 10 Biomass-based resistance and resilience in Amazon forests.
a, The spatial pattern of mean resistance derived from the measured above ground biomass in trees for 5 plots in DS drought in tropical South America, derived from Brienen et al., 2015 (ref 84,85). b, The spatial pattern of mean resistance derived from the measured aboveground biomass in trees for 18 plots in WS drought in tropical South America. c, The spatial pattern of mean resistance derived from the measured aboveground biomass in trees for 15 plots in DS+WS drought in tropical South America. (d-f) The same analysis as (a-c) but for biomass-based resilience in (d) DS, (e) WS, and (f) DS+WS drought. g, Mean ln-transformed resistance (ln(Rt)) for plots that experienced DS, WS, and DS+WS droughts in tropical South America. (h) Mean ln-transformed resilience (ln(Rs)) for plots experienced DS, WS, and DS+WS droughts in tropical South America. Boxplots in (g) and (h) show the median (horizontal lines), mean (triangle), 25th, 40th, 60th, and 75th percentiles (i.e. Q0.25, Q0.4, Q0.6, Q0.75; boxes), and maximum and minimum values (i.e. Max and Min; the top and bottom of the whiskers). Significant differences (p < 0.05) among DS, WS, and DS+WS droughts are denoted by different letters for multiple comparisons. The numbers in the bottom of the panels in (g) and (h) represent the magnitude of plots that experienced DS, WS, and DS+WS droughts, respectively.
Supplementary information
Supplementary information
Supplementary Figs. 1â3 and Tables 1 and 2.
Supplementary Table 3
The metadata of 2,935âtree ring sites used in the study, and extrinsic and intrinsic variables related to drought resistance and resilience obtained from publically available sources. The detailed information (identification, sources and references) of the tree ring data and extrinsic and intrinsic variables are listed under ReadMe.
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Li, X., Piao, S., Wang, K. et al. Temporal trade-off between gymnosperm resistance and resilience increases forest sensitivity to extreme drought. Nat Ecol Evol 4, 1075â1083 (2020). https://doi.org/10.1038/s41559-020-1217-3
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DOI: https://doi.org/10.1038/s41559-020-1217-3