REFERENCES
Allan, E., Weisser, W., Weigelt, A.,
Roscher, C., Fischer, M. & Hillebrand, H. (2011). More diverse plant
communities have higher functioning over time due to turnover in
complementary dominant species. Proc. Natl. Acad. Sci. U. S. A. ,
108, 17034-17039.
Allison, G. (2004). The influence of
species diversity and stress intensity on community resistance and
resilience. Ecological Monographs , 74, 117-134.
Allison, S.D. & Martiny, J.B.H.
(2008). Resistance, resilience, and redundancy in microbial communities.Proc. Natl. Acad. Sci. U. S. A. , 105, 11512-11519.
Anja, V., Michael, S.L. & Alexandra,
W. (2012). Grassland Resistance and Resilience after Drought Depends on
Management Intensity and Species Richness. Plos One , 7, e36992.
Baert, J.M., De Laender, F., Sabbe, K.
& Janssen, C.R. (2016). Biodiversity increases functional and
compositional resistance, but decreases resilience in phytoplankton
communities. Ecology , 97, 3433-3440.
Bai, W., Sun, X., Wang, Z. & Li, L.
(2009). Nitrogen addition and rhizome severing modify clonal growth and
reproductive modes of Leymus chinensis population. Plant Ecology ,
205, 13-21.
Bowman, W.D., Ayyad, A., de Mesquita,
C.P.B., Fierer, N., Potter, T.S. & Sternagel, S. (2018). Limited
ecosystem recovery from simulated chronic nitrogen deposition.Ecological Applications , 28, 1762-1772.
Bray, J.R. & Curtis, J.T. (1957). An
Ordination of the Upland Forest Communities of Southern Wisconsin.Ecological Monographs , 27, 326-349.
Breshears, D.D., Cobb, N.S., Rich,
P.M., Price, K.P., Allen, C.D., Balice, R.G. et al. (2005). Regional
vegetation die-off in response to global-change-type drought.Proc. Natl. Acad. Sci. U. S. A. , 102, 15144-15148.
Buckland, S.M., Grime, J.P., Hodgson,
J.G. & Thompson, K. (1997). A comparison of plant responses to the
extreme drought of 1995 in northern England. Journal of Ecology ,
85, 875-882.
Caldeira, M.C., Hector, A., Loreau,
M. & Pereira, J.S. (2005). Species richness, temporal variability and
resistance of biomass production in a Mediterranean grassland.Oikos , 110, 115-123.
Carter, D.L. & Blair, J.M. (2012).
High richness and dense seeding enhance grassland restoration
establishment but have little effect on drought response.Ecological Applications , 22, 1308-1319.
Ciais, P., Reichstein, M., Viovy, N.,
Granier, A., Ogée, J., Allard, V. et al. (2005). Europe-wide reduction
in primary productivity caused by the heat and drought in 2003.Nature , 437, 529-533.
Clark, D.L. & Wilson, M.V. (2001).
Fire, mowing, and hand-removal of woody species in restoring a native
wetland prairie in the Willamette Valley of Oregon. Wetlands , 21,
135-144.
Cui, Q.A., Lu, X.T., Wang, Q.B. &
Han, X.G. (2010). Nitrogen fertilization and fire act independently on
foliar stoichiometry in a temperate steppe. Plant and Soil , 334,
209-219.
da Costa, A.C.L., Galbraith, D.,
Almeida, S., Portela, B.T.T., da Costa, M., de Athaydes Silva Junior, J.
et al. (2010). Effect of 7 yr of experimental drought on vegetation
dynamics and biomass storage of an eastern Amazonian rainforest.New Phytol. , 187, 579-591.
De Boeck, H.J., Dreesen, F.E.,
Janssens, I.A. & Nijs, I. (2011). Whole‐system responses of
experimental plant communities to climate extremes imposed in different
seasons. New Phytol. , 189, 806-817.
De Boeck, H.J., Lemmens, C.M.H.M.,
Zavalloni, C., Gielen, B., Malchair, S., Carnol, M. et al. (2008).
Biomass production in experimental grasslands of different species
richness during three years of climate warming. Biogeosciences ,
5, 585-594.
DeClerck, F.A.J., Barbour, M.G. &
Sawyer, J.O. (2006). Species richness and stand stability in conifer
forests of the Sierra Nevada. Ecology , 87, 2787-2799.
Dreesen, F.E., Boeck, H.D., Janssens,
I.A. & Nijs, I. (2014). Do successive climate extremes weaken the
resistance of plant communities? An experimental study using plant
assemblages. Biogeosciences , 11, 109-121.
Florence, V., Karim, B. & Mark, N.
(2014). Designing resilient and sustainable grasslands for a drier
future: Adaptive strategies, functional traits and biotic interactions.European Journal of Agronomy , 52, 81-89.
Frank, D.A. & McNaughton, S.J.
(1991). Stability Increases with Diversity in Plant Communities:
Empirical Evidence from the 1988 Yellowstone Drought. Oikos , 62,
360-362.
Friedrich, U., von Oheimb, G.,
Kriebitzsch, W.-U., Schlesselmann, K., Weber, M.S. & Haerdtle, W.
(2012). Nitrogen deposition increases susceptibility to drought -
experimental evidence with the perennial grass Molinia caerulea (L.)
Moench. Plant and Soil , 353, 59-71.
Galvanek, D., Mutnanova, M. & Dite,
D. (2015). Restoration mowing of a calcareous fen - response of species
to re-applied management measures. Biologia , 70, 349-355.
Grime, J.P. (1998). Benefits of plant
diversity to ecosystems: immediate, filter and founder effects.Journal of Ecology , 86, 902-910.
Grime, J.P., Brown, V.K., Thompson,
K., Masters, G.J., Hillier, S.H., Clarke, I.P. et al. (2000). The
response of two contrasting limestone grasslands to simulated climate
change. Science , 289, 762-765.
Grman, E., Lau, J.A., Schoolmaster,
D.R. & Gross, K.L. (2010). Mechanisms contributing to stability in
ecosystem function depend on the environmental context. Ecology
Letters , 13, 1400-1410.
Guelzow, N., Muijsers, F., Ptacnik,
R. & Hillebrand, H. (2017). Functional and structural stability are
linked in phytoplankton metacommunities of different connectivity.Ecography , 40, 719-732.
Hautier, Y., Seabloom, E.W., Borer,
E.T., Adler, P.B., Harpole, W.S., Hillebrand, H. et al. (2014).
Eutrophication weakens stabilizing effects of diversity in natural
grasslands. Nature , 508, 521-525.
Hautier, Y., Tilman, D., Isbell, F.,
Seabloom, E.W., Borer, E.T. & Reich, P.B. (2015). Anthropogenic
environmental changes affect ecosystem stability via biodiversity.Science , 348, 336-340.
Hautier, Y., Zhang, P., Loreau, M.,
Wilcox, K.R., Seabloom, E.W., Borer, E.T. et al. (2020). General
destabilizing effects of eutrophication on grassland productivity at
multiple spatial scales. Nature Communications , 11, 5375.
Hector, A., Hautier, Y., Saner, P.,
Wacker, L., Bagchi, R., Joshi, J. et al. (2010). General stabilizing
effects of plant diversity on grassland productivity through population
asynchrony and overyielding. Ecology , 91, 2213-2220.
Heitschmidt, R.K., Klement, K.D. &
Haferkamp, M.R. (2005). Interactive Effects of Drought and Grazing on
Northern Great Plains Rangelands. Rangeland Ecology &
Management , 58, 11-19.
Hillebrand, H. & Kunze, C. (2020).
Meta-analysis on pulse disturbances reveals differences in functional
and compositional recovery across ecosystems. Ecology Letters ,
23, 575-585.
Hillebrand, H., Langenheder, S.,
Lebret, K., Lindstrom, E., Oestman, O. & Striebel, M. (2018).
Decomposing multiple dimensions of stability in global change
experiments. Ecology Letters , 21, 21-30.
Hofer, D., Suter, M., Buchmann, N. &
Luscher, A. (2017). Nitrogen status of functionally different forage
species explains resistance to severe drought and post-drought
overcompensation. Agric. Ecosyst. Environ. , 236, 312-322.
Hoover, D.L., Knapp, A.K. & Smith,
M.D. (2014). Resistance and resilience of a grassland ecosystem to
climate extremes. Ecology , 95, 2646-2656.
IPCC (2014). Climate Change 2014:
Synthesis Report. Contribution of Working Groups I, II and III to the
Fifth Assessment. IPCC, Geneva, Switzerland.
Isbell, F., Craven, D., Connolly, J.,
Loreau, M., Schmid, B., Beierkuhnlein, C. et al. (2015). Biodiversity
increases the resistance of ecosystem productivity to climate extremes.Nature , 526, 574-577.
Isbell, F.I., Polley, H.W. & Wilsey,
B.J. (2009). Biodiversity, productivity and the temporal stability of
productivity: patterns and processes. Ecology Letters , 12,
443-451.
Ives, A.R. & Carpenter, S.R. (2007).
Stability and Diversity of Ecosystems. Science , 317, 58-62.
Ives, A.R., Gross, K. & Klug, J.L.
(1999). Stability and Variability in Competitive Communities.Science , 286, 542-544.
Jasper, V.R. & Frank, B. (2010).
Diversity enhances community recovery, but not resistance, after
drought. Journal of Ecology , 98, 81–86.
Kahmen, A., Perner, J. & Buchmann,
N. (2005). Diversity-dependent productivity in semi-natural grasslands
following climate perturbations. Funct. Ecol. , 19, 594-601.
Kaye, J.P. & Hart, S.C. (1998).
Ecological Restoration Alters Nitrogen Transformations in a Ponderosa
Pine-Bunchgrass Ecosystem. Ecological Applications , 8, 1052-1060.
Kinugasa, T., Tsunekawa, A. &
Shinoda, M. (2012). Increasing nitrogen deposition enhances post-drought
recovery of grassland productivity in the Mongolian steppe.Oecologia , 170, 857-865.
Knapp, A.K., Carroll, C.J.W., Denton,
E.M., Pierre, K.J.L., Collins, S.L. & Smith, M.D. (2015). Differential
sensitivity to regional-scale drought in six central US grasslands.Oecologia , 177, 949-957.
Kreyling, J., Dengler, J., Walter,
J., Velev, N., Ugurlu, E., Sopotlieva, D. et al. (2017). Species
richness effects on grassland recovery from drought depend on community
productivity in a multisite experiment. Ecology letters , 20,
1405-1413.
Leps, J. (2014). Scale- and
time-dependent effects of fertilization, mowing and dominant removal on
a grassland community during a 15-year experiment. Journal of
Applied Ecology , 51, 978-987.
Liu, X., Duan, L., Mo, J., Du, E.,
Shen, J., Lu, X. et al. (2011). Nitrogen deposition and its ecological
impact in China: an overview. Environmental Pollution , 159,
2251-2264.
Loreau, M. & de Mazancourt, C.
(2008). Species synchrony and its drivers: Neutral and nonneutral
community dynamics in fluctuating environments. Am. Nat. , 172,
E48-E66.
Lu, X., Rao, X. & Dong, W. (2021).
Model evaluation and uncertainties in projected changes of drought over
northern China based on CMIP5 models. International Journal of
Climatology , 41, E3085-E3100.
Macgillivray, C.W., Grime, J.P.,
Band, S.R., Booth, R.E., Campbell, B., Hendry, G.A.F. et al. (1995).
Testing predictions of the resistance and resilience of vegetation
subjected to extreme events. Funct. Ecol. , 9, 640-649.
Mackie, K.A., Zeiter, M., Bloor,
J.M.G. & Stampfli, A. (2019). Plant functional groups mediate drought
resistance and recovery in a multisite grassland experiment.Journal of Ecology , 107, 937-949.
Maron, J.L. & Jefferies, R.L.
(2001). Restoring enriched grasslands: effects of mowing on species
richness, productivity, and nitrogen retention. Ecological
Applications , 11, 1088-1100.
May, R.M. (1973). Stability and
complexity in model ecosystems. Monographs in population biology ,
6, 1-235.
Meehl, G.A., Karl, T., Easterling,
D.R., Changnon, S., Pielke Jr, R., Changnon, D. et al. (2000). An
introduction to trends in extreme weather and climate events:
observations, socioeconomic impacts, terrestrial ecological impacts, and
model projections. Bulletin of the American Meteorological
Society , 81, 413-416.
Molina, C.D., Tognetti, P.M., Graff,
P. & Chaneton, E.J. (2021). Mowing does not redress the negative effect
of nutrient addition on alpha and beta diversity in a temperate
grassland. Journal of Ecology , 109, 1501-1510.
Pan, Q.-M., Bai, Y.-F., Han, X.-G. &
Yang, J.-C. (2005). Effects of nitrogen additions on a Leymus chinensis
population in typical steppe of inner Mongolia. Zhiwu Shengtai
Xuebao , 29, 311-317.
Peñuelas, J., Prieto, P., Beier, C.,
Cesaraccio, C., De Angelis, P., De Dato, G. et al. (2007). Response of
plant species richness and primary productivity in shrublands along a
north–south gradient in Europe to seven years of experimental warming
and drought: reductions in primary productivity in the heat and drought
year of 2003. Global Change Biology , 13, 2563-2581.
Pfisterer, A.B. & Schmid, B. (2002).
Diversity-dependent production can decrease the stability of ecosystem
functioning. Nature , 416, 84-86.
Polazzo, F. & Rico, A. (2021).
Effects of multiple stressors on the dimensionality of ecological
stability. Ecology Letters , DOI: 10.1111/ele.13770, 1-13.
Ruppert, J.C., Harmoney, K., Henkin,
Z., Snyman, H.A., Sternberg, M., Willms, W. et al. (2015). Quantifying
drylands’ drought resistance and recovery: the importance of drought
intensity, dominant life history and grazing regime. Global Change
Biology , 21, 1258-1270.
Shi, B., Wang, Y., Meng, B., Zhong,
S. & Sun, W. (2018). Effects of Nitrogen Addition on the Drought
Susceptibility of the Leymus chinensis Meadow Ecosystem Vary with
Drought Duration. Frontiers in Plant Science , 9, 254.
Shurin, J.B., Arnott, S.E.,
Hillebrand, H., Longmuir, A., Pinel-Alloul, B., Winder, M. et al.
(2007). Diversity–stability relationship varies with latitude in
zooplankton. Ecology Letters , 10, 127-134.
Socher, S.A., Prati, D., Boch, S.,
Müller, J., Baumbach, H., Gockel, S. et al. (2013). Interacting effects
of fertilization, mowing and grazing on plant species diversity of 1500
grasslands in Germany differ between regions. Basic and Applied
Ecology , 14, 126-136.
Song, M.-H. & Yu, F.-H. (2015).
Reduced compensatory effects explain the nitrogen-mediated reduction in
stability of an alpine meadow on the Tibetan Plateau. New
Phytol. , 207, 70-77.
Stampfli, A., Bloor, J.M.G., Fischer,
M. & Zeiter, M. (2018). High land-use intensity exacerbates shifts in
grassland vegetation composition after severe experimental drought.Glob Chang Biol , 24, 2021-2034.
Suding, K.N., Collins, S.L., Gough,
L., Clark, C., Cleland, E.E., Gross, K.L. et al. (2005). Functional- and
abundance-based mechanisms explain diversity loss due to N
fertilization. Proc. Natl. Acad. Sci. U. S. A. , 102, 4387-4392.
Tian, Q., Liu, N., Bai, W., Li, L.,
Chen, J., Reich, P.B. et al. (2016). A novel soil manganese mechanism
drives plant species loss with increased nitrogen deposition in a
temperate steppe. Ecology , 97, 65-74.
Tilman, D. & Downing, J.A. (1994).
Biodiversity and stability in grasslands. Nature , 367, 363-365.
Valliere, J.M., Irvine, I.C.,
Santiago, L. & Allen, E.B. (2017). High N, dry: Experimental nitrogen
deposition exacerbates native shrub loss and nonnative plant invasion
during extreme drought. Global Change Biology , 23, 4333-4345.
Van Peer, L., Nijs, I., Reheul, D. &
De Cauwer, B. (2004). Species richness and susceptibility to heat and
drought extremes in synthesized grassland ecosystems: compositional vs
physiological effects. Funct. Ecol. , 18, 769-778.
Van Ruijven, J. & Berendse, F.
(2010). Diversity enhances community recovery, but not resistance, after
drought. Journal of Ecology , 98, 81-86.
Vitousek, P.M., Aber, J.D., Howarth,
R.W., Likens, G.E., Matson, P.A., Schindler, D.W. et al. (1997). Human
alteration of the global nitrogen cycle: sources and consequences.Ecological applications , 7, 737-750.
Vogel, A., Scherer-Lorenzen, M. &
Weigelt, A. (2012). Grassland Resistance and Resilience after Drought
Depends on Management Intensity and Species Richness. PLoS ONE ,
7, e36992.
Wagg, C., O’Brien, M.J., Vogel, A.,
Scherer-Lorenzen, M., Eisenhauer, N., Schmid, B. et al. (2017). Plant
diversity maintains long-term ecosystem productivity under frequent
drought by increasing short-term variation. Ecology , 98,
2952-2961.
Wang, S., Wilkes, A., Zhang, Z.,
Chang, X., Lang, R., Wang, Y. et al. (2011). Management and land use
change effects on soil carbon in northern China’s grasslands: a
synthesis. Agriculture, ecosystems & environment , 142, 329-340.
Wang, Y.F., Yu, S.X. & Wang, J.
(2007). Biomass-dependent susceptibility to drought in experimental
grassland communities. Ecology Letters , 10, 401-410.
Wardle, D.A., Bonner, K.I. & Barker,
G.M. (2000). Stability of ecosystem properties in response to
above-ground functional group richness and composition. Oikos ,
89, 11-23.
Williams, D.W., Jackson, L.L. &
Smith, D.D. (2010). Effects of Frequent Mowing on Survival and
Persistence of Forbs Seeded into a Species-Poor Grassland. Restor.
Ecol. , 15, 24-33.
Xu, Q., Yang, X., Yan, Y., Wang, S.,
Loreau, M. & Jiang, L. (2021). Consistently positive effect of species
diversity on ecosystem, but not population, temporal stability.Ecology Letters , DOI: 10.1111/ele.13777.
Xu, Z., Ren, H., Cai, J., Wang, R.,
He, P., Li, M.-H. et al. (2015a). Antithetical effects of nitrogen and
water availability on community similarity of semiarid grasslands:
evidence from a nine-year manipulation experiment. Plant and
Soil , 397, 357-369.
Xu, Z., Ren, H., Cai, J., Wang, R.,
Li, M.H., Wan, S. et al. (2014). Effects of experimentally-enhanced
precipitation and nitrogen on resistance, recovery and resilience of a
semi-arid grassland after drought. Oecologia , 176, 1187-1197.
Xu, Z., Ren, H., Li, M.-H., van
Ruijven, J., Han, X., Wan, S. et al. (2015b). Environmental changes
drive the temporal stability of semi-arid natural grasslands through
altering species asynchrony. Journal of Ecology , 103, 1308-1316.
Xu, Z.Z. & Zhou, G.S. (2006).
Nitrogen metabolism and photosynthesis in Leymus chinensis in response
to long-term soil drought. Journal of Plant Growth Regulation ,
25, 252-266.
Yachi, S. & Loreau, M. (1999).
Biodiversity and ecosystem productivity in a fluctuating environment:
The insurance hypothesis. Proc. Natl. Acad. Sci. U. S. A. , 96,
1463-1468.
Yang, G.-J., Lu, X.-T., Stevens,
C.J., Zhang, G.-M., Wang, H.-Y., Wang, Z.-W. et al. (2019). Mowing
mitigates the negative impacts of N addition on plant species diversity.Oecologia , 189, 769-779.
Yang, H., Jiang, L., Li, L., Li, A.,
Wu, M. & Wan, S. (2012). Diversity-dependent stability under mowing and
nutrient addition: evidence from a 7-year grassland experiment.Ecology Letters , 15, 619-626.
Yang, Y.F., Liu, G.C. & Zhang, B.T.
(1995). An analysis of age structure and the strategy for asexual
propagation of Aneurolepidium Chinense population. Chinese
Bulletin of Botany , 37, 147-153.
Zhang, Y., Loreau, M., He, N., Zhang,
G. & Han, X. (2017). Mowing exacerbates the loss of ecosystem stability
under nitrogen enrichment in a temperate grassland. Funct. Ecol. ,
31, 1637-1646.
Zhang, Y., Loreau, M., Lu, X., He,
N., Zhang, G. & Han, X. (2016). Nitrogen enrichment weakens ecosystem
stability through decreased species asynchrony and population stability
in a temperate grassland. Global Change Biology , 22, 1445-1455.
Zhang, Y.H., Feng, J.C., Loreau, M.,
He, N.P., Han, X.G. & Jiang, L. (2019). Nitrogen addition does not
reduce the role of spatial asynchrony in stabilising grassland
communities. Ecology Letters , 22, 563-571.
Table 1 Results
(F -value) of linear mixed-effects models on the effects of block
(B), nitrogen addition (N), mowing (M) and their interactions on soil
pH, soil inorganic nitrogen concentration (soil IN), community
aboveground biomass (AGB), species richness, asynchrony, biomass
resistance (B rst), biomass recovery (B rc), structural
resistance (S rst), structural recovery (S rc) of plant
communities, B rst and B rc of L. chinensis ,S rst and S rc of dominant species experienced drought in
three consecutive growing seasons. ***p < 0.001,
**p < 0.01, *p < 0.05, ^ p< 0.1, ns p > 0.1.