Acta Oecologica 98 (2019) 25–29

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Acta Oecologica
journal homepage: www.elsevier.com/locate/actoec

Alpine grassland degradation reduced plant species diversity and stability of T

plant communities in the Northern Tibet Plateau
Congyan Wang∗, Mei Wei, Bingde Wu, Shu Wang, Kun Jiang
Institute of Environment and Ecology, Academy of Environmental Health and Ecological Security & School of the Environment and Safety Engineering, Jiangsu University,
Zhenjiang, 212013, PR China

A R T I C LE I N FO A B S T R A C T

Keywords: The alpine grassland in the Northern Tibet Plateau has been degraded in recent decades. This study aimed at
Alpine grassland addressing the effects of the degree of degradation on species diversity and stability of plant communities and
Community stability the relationships between diversity stability. Stability of plant communities decreased with the increasing degree
Degradation of degradation probably due to the significantly decreased diversity and evenness of plant communities with
Diversity-stability correlation
degradation. Diversity (including Shannon and Simpson diversity, Pielou evenness, and species richness) was
Northern Tibet Plateau
Plant species diversity
positively correlated with the corresponding stability of plant communities. This suggets that high levels of plant
diversity lead to higher levels of complementary resource use via niche complementation. The relative abun-
dance of the most dominant species was negatively correlated with the stability of the corresponding plant
communities. This suggests that the stability of plant communities decreases with increasing degradation but the
relative abundance of the most dominant species increases with degradation. In conclusion, diversity and the
corresponding stability of plant communities decrease with increasing degradation in the alpine grassland in the
Northern Tibet Plateau.

1. Introduction economic conditions as well as the environmental health and ecological

security (Shao and Cai, 2008; Gao et al., 2010; Wen et al., 2010; Zhao
Grassland ecosystems are the largest terrestrial ecosystems in the et al., 2013; Rong et al., 2018). One important signs of degradation in
world, accounting for about half of the world's land area, and play an the alpine grassland in the Northern Tibet Plateau is the reduction of
extremely important role in the stability of global ecosystem func- diversity of plant communities (Han et al., 2008; Shao and Cai, 2008;
tioning (Reeder and Schuman, 2002). The Northern Tibet Plateau Gao et al., 2010; Zhao et al., 2013). Diversity of plant communities is
boasts vast natural grassland resources, with various types of natural closely associated with their stability (Tilman and Downing, 1994;
grasslands covering 4.8 × 106 hm2, accounting for 81% of the land area Grime et al., 2000; Báez and Collins, 2008) via the enhanced ecosystem
of the Northern Tibet Plateau and 59% of the natural grassland area of functioning and the sustained ecosystem services (Gamfeldt et al.,
the Tibet Autonomous Region (Hu, 2000). Thus, the Northern Tibet 2008; Zavaleta et al., 2010; Isbell et al., 2011). Consequently, it is
Plateau is the main pastoral area in Tibet Autonomous Region (Zhao important to determine the effects of degradation on plant diversity and
et al., 2013; Baima, 2016). However, due to the particularity of the the corresponding stability of communities in the alpine grassland in
climate and geographical location, the Northern Tibet Plateau possesses the Northern Tibet Plateau.
unique ecological characteristics, namely, simple structure, poor repair There are two conflicting hypotheses about community stability.
ability, weak stability and easiness to be disturbed by external factors According to the diversity hypothesis, the temporal stability of plant
(Gao et al., 2010; Baima, 2016; Rong et al., 2018). Unfortunately, the communities is positively correlated with plant species diversity, i.e.,
alpine grassland in the Northern Tibet Plateau is experiecing degrada- greater levels of diversity stabilize community fluctuations via the
tion processes due to the effects of both anthropogenic and natural niche complementation mainly by the more efficient complementary
factors (such as over grazing, urban construction, road construction, resource use (Cottingham et al., 2001; Ives and Carpenter, 2007; De
climate warming, etc.). The degradation in the alpine grassland can Mazancourt et al., 2013; Loreau and De Mazancourt, 2013). This hy-
seriously affect the sustainable development of local social and pothesis has been confirmed in numerous studies reporting a

∗
Corresponding author. Institute of Environment and Ecology, Academy of Environmental Health and Ecological Security & School of the Environment and Safety
Engineering, Jiangsu University, No. 301, Xuefu Road, Zhenjiang, 212013, PR China.
E-mail address: [email protected] (C. Wang).

https://doi.org/10.1016/j.actao.2019.05.005
Received 25 September 2018; Received in revised form 11 May 2019; Accepted 26 May 2019
Available online 28 May 2019
1146-609X/ © 2019 Elsevier Masson SAS. All rights reserved.
C. Wang, et al. Acta Oecologica 98 (2019) 25–29

significantly higher stability in more diverse communities than in the sampling area is entierely flat, which eliminate the possible effects of
less diverse ones (Isbell et al., 2011; Cardinale et al., 2012; Gross et al., differences in terrain. The sampling area including three sampling sites
2014; Tilman et al., 2014). Conversely, the mass ratio hypothesis pro- with different degrees of degradation. The degree of degradation was
poses that the stability of plant communities is positively correlated determined based on the plant cover and divided into one of three le-
with the stability of the most dominant species (the species with the vels for each sampling quadrat: undegraded (> 75%), moderately de-
highest relative abundance), i.e., the most dominant species are the graded (50%–75%), and heavy degraded (< 35%). In particular,
foremost determining factor driving the stability of plant communities moderately degraded conditions mimicked the initial stage of the
(Grime, 1998; Hillebrand et al., 2008; Valone and Balaban-Feld, 2018). grassland degradation; heavy degraded conditions mimicked the late
Numerous studies have also verified this hypothesis (Sasaki and stage of the grassland degradation; and undegraded conditions mi-
Lauenroth, 2011; Hallett et al., 2014; Yang et al., 2017). Therefore, it is micked the basically original grassland without obvious degradation.
doubtful whether the reduction of plant diversity in the alpine grass- The three sampling sites were more than 500 m apart. Thirty replicate
land in the Northern Tibet Plateau mediated by the degradation will quadrats (1 m × 1 m) were evaluated in each of the three sampling
lead to the reduction of the corresponding stability of communities. sites. Sampling quadrats were placed at a minimum distance of 10 m
The present study aims to give insights into the effects of the de- from each to other. All quadrats were checked to determine the number
gradation on plant species diversity and community stability in the of individuals per plant species, the number of individuals of all plant
alpine grassland in the Northern Tibet Plateau. species, and the number of plant species. The relative abundance of
In particular, this study tests the following hypotheses: (1) Diversity each particular species was evaluated using the ratio of the number of
and the corresponding stability of plant communities in the alpine individuals of this species to the number of individuals of all plant
grassland in the Northern Tibet Plateau decreases markedly with in- species in the measured quadrats (Guiasu and Guiasu, 2012; Melese and
creasing degree of degradation; and (2) diversity of plant communities Ayele, 2017; Wang et al., 2018a, b).
is positively correlated with the corresponding stability of plant com-
munities.
2.2. Methods for determining the diversity and stability of plant
communities
2. Materials and methods
The diversity of plant communities in the degraded alpine grassland
2.1. Experimental design in the Northern Tibet Plateau was quantified using Shannon's diversity
index (H′), Simpson's diversity index (D), Pielou's evenness index (EH),
In early-June 2018, sampling quadrats of alpine grassland vegeta- and Margalef's richness index (F). Shannon's diversity index was cal-
tion with different degrees of degradation were randomly selected in culated as H’ = −ΣPilnPi (Shannon and Weaver, 1949), where Pi is the
the Northern Tibet Plateau in Shenza, Nagqu, Tibet (30°56′46″ N, relative abundance of plant species i in one particular quadrat. Pi was
88°37′54″ E), in a area with a semi-arid climate of the plateau subfrigid defined as Pi = ni/N, where ni is the number of individuals of plant
zone located at about 4700 m above sea level. According to local cli- species i and N is the number of individuals of all plant species in one
mate records (Wang, 2015), the annual mean temperature at the area is particular quadrat. Simpson's diversity index was calculated as
approximately 0.2 °C with a maximum of 25.1 °C and a minimum of D = 1−Σ(ni/N)2 (Simpson, 1949). Pielou's evenness index was calcu-
−30.1 °C. The annual precipitation is approximately 299 mm. Fig. 1 lated as EH = H'/lnS (Pielou, 1966), where S is the number of plant
shows the geographic location of the sampling area. species in one particular quadrat. Margalef's richness index was calcu-
The sampling area is composed of alpine grassland communities. lated as F = (S–1)/lnN (Margalef, 1951).
There were no shrubs or trees in any sampling communities surveyed in The stability of plant communities in the degraded alpine grassland
this study. The sampling area is devoid of trees and shrubs due to the in the Northern Tibet Plateau was determined by using inverse of
low average annual temperature and precipitation. The selected coefficient of variation ICV. ICV = μ/σ, where μ is the average relative
abundance of all plant species in one particular quadrat and σ is the
standard deviation for the average relative abundance of all plant
species in one particular quadrat (Sasaki and Lauenroth, 2011; Yang
et al., 2011; Valone and Balaban-Feld, 2018).

2.3. Statistical analysis

Deviations from normality and homogeneity of the variances were

determined before data analysis. Differences in the values for the in-
dices of plant species diversity and the stability of corresponding plant
communities in the alpine grassland in the Northern Tibet Plateau with
different degrees of degradation were evaluated by analysis of variance
followed by Student–Newman–Keuls test for multiple comparisons.
Correlation analyses were performed using the Spearman's rank corre-
lation coefficient to determine the relationships between the relative
abundance of the most dominant species and community stability index
as well as the relationships among the indices of plant species diversity
and community stability index. A probability value equal to or less than
0.05 was used to determine statistical significance. All statistical ana-
lyses were performed using IBM SPSS Statistics (version 25.0; IBM
Corp., Armonk, NY, USA).
Fig. 1. Approximate geographic location of the sampling area (blue triangle).
(For interpretation of the references to colour in this figure legend, the reader is
referred to the Web version of this article.)

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C. Wang, et al. Acta Oecologica 98 (2019) 25–29

Fig. 2. Differences in the relative abundance of the most dominant species and
the indices of diversity and community stability of plant communities in the
degraded alpine grassland in the Northern Tibet Plateau. Bars (mean with
standard error, n = 30) with different lowercase letters indicate a significant
difference (P < 0.05) assessed by one-way analysis of variance followed by
Student–Newman–Keuls tests for multiple comparisons. “ns” means no sig-
nificant difference (P > 0.05).

3. Results

3.1. Differences in plant diversity and stability of communities

Differences in diversity and stability of plant communities were

detected across different degrees of degradation in the alpine grassland
in the Northern Tibet Plateau (Fig. 2). In particular, Shannon's diversity
index, Simpson's diversity index, and Pielou's evenness index of plant
communities significantly decreased with increasing degree of de-
gradation (P < 0.05; Fig. 2a–c). Margalef's richness index and com-
munity stability index also decreased with increasing degree of de-
gradation, although this change was not significant (P > 0.05; Fig. 2d
and e). However, the relative abundance of the most dominant species
significantly increased with increasing degree of degradation
(P < 0.05; Fig. 2f).

3.2. Relationships between relative abundance of the most dominant species

and community stability of plant communities

The relative abundance of the most dominant species was negatively

correlated with community stability of plant communities for all de-
grees of degradation (P < 0.01; Table 1).
Shannon's diversity index, Simpson's diversity index, and Pielou's
evenness index of plant communities were positively correlated with
stability (P < 0.0001; Table 2). Margalef's richness index was also
positively correlated with stability but did not reach statistical sig-
nificance (P > 0.05; Table 2).

4. Discussion

Plant communities may reveal temporal fluctuations in abundance

and/or biomass as time goes on (Davis et al., 2000; Valone and
Balaban-Feld, 2018; Wang et al., 2018a, b). It is important to take into
consideration the effects of shifts in the structure and composition of
plant communities on the corresponding stability because these
changes can exert powerful effects on the capabilities of ecosystem
services (De Mazancourt et al., 2013; Gross et al., 2014; Tilman et al.,
2014). Our study indicates that the stability of plant communities de-
creased with increasing degree of degradation in the alpine grassland in
the Northern Tibet Plateau. This implies that the degradation in the
alpine grassland in the Northern Tibet Plateau can significantly reduce
the capabilities of ecosystem services. The phenomenon is most

Table 1
Relationships between the relative abundance of the most dominant species and
community stability index of plant communities in the alpine grassland in the
Northern Tibet Plateau with different degrees of degradation (Spearman's rank
correlation coefficient, ρ). ** and *** indicate significant differences at 0.01 and
0.001 probability levels, respectively. P values equal to or less than 0.05 are
shown in bold.
The degree of degradation Values

Undegraded ρ −0.469**
P 0.0089
Moderately degraded ρ −0.880***
P < 0.0001
Heavy degraded ρ −0.860***
P < 0.0001

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Table 2 Author contribution

Relationships between diversity and stability of plant communities in the alpine
grassland in Northern Tibet Plateau with different degrees of degradation Congyan Wang conceived and designed this study. Mei Wei and
(Spearman's rank correlation coefficient, ρ). *** indicate significant differences Bingde Wu performed the experiments. Shu Wang and Kun Jiang ana-
at 0.001 probability levels. P values equal to or less than 0.05 are shown in
lyzed the data. Mei Wei and Bingde Wu contributed reagents/mate-
bold.
rials/analysis tools. Congyan Wang wrote the manuscript. All authors
The indices of plant species diversity Values have read the manuscript and have agreed to submit it in its current
form for consideration for publication.
Shannon's diversity index ρ 0.597***
P < 0.0001
Simpson's diversity index ρ 0.689*** Acknowledgements
P < 0.0001
Pielou's evenness index ρ 0.762*** This study was supported by National Key Research and
P < 0.0001
Development Program of China (2016YFC0502002), National Natural
Margalef's richness index ρ 0.180
P 0.0903 Science Foundation of China (31300343), and Jiangsu Collaborative
Innovation Center of Technology and Material of Water Treatment. We
are very grateful to the anonymous reviewers for the insightful and
probably caused by the significantly decreased diversity and evenness constructive comments that greatly improved this manuscript.
of plant communities with increasing degree of degradation because
high levels of plant species diversity can minimize community fluc- References
tuations through the ecological niche complementation, mainly by the
greater levels of complementary resource use (Cottingham et al., 2001; Báez, S., Collins, S.L., 2008. Shrub invasion decreases diversity and alters community
Ives and Carpenter, 2007; De Mazancourt et al., 2013; Loreau and De stability in Northern Chihuahuan Desert plant communities. PLoS One 3, e2332.
Baima, Y.Z., 2016. The present situation of grassland resources in Nagqu region of Tibet
Mazancourt, 2013). We also found a strong correlation between the and its development and utilization countermeasures. South China Agr. 10, 119–120
diversity and evenness and the corresponding stability of plant com- (In Chinese).
munities. This finding was consistent with our first hypothesis. How- Byun, C., De Blois, S., Brisson, J., 2013. Plant functional group identity and diversity
determine biotic resistance to invasion by an exotic grass. J. Ecol. 101, 128–139.
ever, the result of this study showed no significant correlation between Cardinale, B.J., Duffy, J.E., Gonzalez, A., Hooper, D.U., Perrings, C., Venail, P., Narwani,
the richness and the corresponding stability of plant communities. Ac- A., Mace, G.M., Tilman, D., Wardle, D.A., Kinzig, A.P., Daily, G.C., Loreau, M., Grace,
cordingly, diversity and evenness, rather than richness, were the most J.B., Larigauderie, A., Srivastava, D.S., Naeem, S., 2012. Biodiversity loss and its
impact on humanity. Nature 486, 59–67.
important drivers of stability of plant communities. Cardinale, B.J., Gross, K., Fritschie, K., Flombaum, P., Fox, J.W., Rixen, C., Wilsey, B.J.,
Plant diversity and evenness were strongly correlated with the 2013. Biodiversity simultaneously enhances the production and stability of commu-
corresponding stability of communities, which suggests that high levels nity biomass, but the effects are independent. Ecology 94, 1697–1707.
Cottingham, K.L., Brown, B.L., Lennon, J.T., 2001. Biodiversity may regulate the temporal
of diversity lead to higher levels of complementary resource use via the
variability of ecological systems. Ecol. Lett. 4, 72–85.
niche complementation (Cottingham et al., 2001; Ives and Carpenter, Davis, M.A., Grime, J.P., Thompson, K., 2000. Fluctuating resources in plant commu-
2007; De Mazancourt et al., 2013; Loreau and De Mazancourt, 2013). nities: a general theory of invasibility. J. Ecol. 88, 528–534.
De Mazancourt, C., Isbell, F., Larocque, A., Berendse, F., De Luca, E., Grace, J.B.,
The underlying mechanisms for the strongly positive diversity-stability
Haegeman, B., Wayne Polley, H., Roscher, C., Schmid, B., Tilman, D., van Ruijven, J.,
correlation may include: (1) complementary effects (multiple plant Weigelt, A., Wilsey, B.J., Loreau, M., 2013. Predicting ecosystem stability from
species can generate more efficient use of ecological resources when community composition and biodiversity. Ecol. Lett. 16, 617–625.
they coexist; Lehman and Tilman, 2000; Isbell et al., 2009; Byun et al., Gamfeldt, L., Hillebrand, H., Jonsson, P.R., 2008. Multiple functions increase the im-
portance of biodiversity for overall ecosystem functioning. Ecology 89, 1223–1231.
2013), (2) species asynchrony effects (the decreases in the abundance of Gao, Q.Z., Duan, M.J., Wan, Y.F., Li, Y.E., Guo, Y.Q., Jiangcun, W.Z., 2010.
some plant species can be neutralized by the increases in the abundance Comprehensive evaluation of eco‒environmental sensitivity in Northern Tibet. Acta
of others; Yang et al., 2012; Loreau and De Mazancourt, 2013; Hautier Ecol. Sin. 30, 4129–4136 (In Chinese).
Gherardi, L.A., Sala, O.E., 2015. Enhanced interannual precipitation variability increases
et al., 2014), (3) portfolio effects (statistical averaging in which tem- plant functional diversity that in turn ameliorates negative impact on productivity.
poral variance in species abundances scales with mean species abun- Ecol. Lett. 18, 1293–1300.
dance; Isbell et al., 2009; Yang et al., 2012; Gherardi and Sala, 2015), Grime, J.P., 1998. Benefits of plant diversity to ecosystems: immediate, filter and founder
effects. J. Ecol. 86, 902–910.
and (4) covariance effects (negative covariance in the abundance of Grime, J.P., Brown, V.K., Thompson, K., Masters, G.J., Hillier, S.H., Clarke, I.P., Askew,
competing species will be increased in the communities with greater A.P., Corker, D., Kielty, J.P., 2000. The response of two contrasting limestone
plant species diversity; Hector et al., 2010; Cardinale et al., 2013). grasslands to simulated climate change. Science 289, 762–765.
Gross, K., Cardinale, B.J., Fox, J.W., Gonzalez, A., Loreau, M., Polley, H.W., Reich, P.B.,
However, we found a significantly negative correlation between the
van Ruijven, J., 2014. Species richness and the temporal stability of biomass pro-
relative abundance of the most dominant species and the corresponding duction: a new analysis of recent biodiversity experiments. Am. Nat. 183, 1–12.
stability of plant communities in the degraded alpine grassland in the Guiasu, R.C., Guiasu, S., 2012. The weighted Gini‒Simpson Index: revitalizing an old
index of biodiversity. Int. J. Ecol. 478728 2012.
Northern Tibet Plateau. The dominant reason can probably be ascribed
Hallett, L.M., Hsu, J.S., Cleland, E.E., Collins, S.L., Dickson, T.L., Farrer, E.C., Gherardi,
to the fact that the stability of plant communities decreased with in- L.A., Gross, K.L., Hobbs, R.J., Turnbull, L., Suding, K.N., 2014. Biotic mechanisms of
creasing degree of degradation, but the relative abundance of the most community stability shift along a precipitation gradient. Ecology 95, 1693–1700.
dominant species increased with increasing degree of degradation. In Han, J.G., Zhang, Y.J., Wang, C.J., Bai, W.M., Wang, Y.R., Han, G.D., Li, L.H., 2008.
Rangeland degradation and restoration management in China. Rangel. J. 30,
particular, the significantly increased relative abundance of the most 233–239.
dominant species can cause the loss of some rare species. Accordingly, Hautier, Y., Seabloom, E.W., Borer, E.T., Adler, P.B., Harpole, W.S., Hillebrand, H., Lind,
the results of this study support the diversity hypothesis (Cottingham E.M., MacDougall, A.S., Stevens, C.J., Bakker, J.D., Buckley, Y.M., Chu, C., Collins,
S.L., Daleo, P., Damschen, E.I., Davies, K.F., Fay, P.A., Firn, J., Gruner, D.S., Jin, V.L.,
et al., 2001; Ives and Carpenter, 2007; De Mazancourt et al., 2013; Klein, J.A., Knops, J.M., La Pierre, K.J., Li, W., McCulley, R.L., Melbourne, B.A.,
Loreau and De Mazancourt, 2013) rather than the mass ratio hypothesis Moore, J.L., O'Halloran, L.R., Prober, S.M., Risch, A.C., Sankaran, M., Schuetz, M.,
(Grime, 1998; Hillebrand et al., 2008; Valone and Balaban-Feld, 2018). Hector, A., 2014. Eutrophication weakens stabilizing effects of diversity in natural
grasslands. Nature 508, 521–525.
The finding was consistent with our second hypothesis. All in all, the Hector, A., Hautier, Y., Saner, P., Wacker, L., Bagchi, R., Joshi, J., Scherer‒Lorenzen, M.,
decreased stability of plant communities in the alpine grassland in the Spehn, E.M., Bazeley‒White, E., Weilenmann, M., Caldeira, M.C., Dimitrakopoulos,
Northern Tibet Plateau under degradation conditions may be due lar- P.G., Finn, J.A., Huss‒Danell, K., Jumpponen, A., Mulder, C.P., Palmborg, C., Pereira,
J.S., Siamantziouras, A.S., Terry, A.C., Troumbis, A.Y., Schmid, B., Loreau, M., 2010.
gely to the decreased diversity and evenness of plant communities.
General stabilizing effects of plant diversity on grassland productivity through po-
pulation asynchrony and overyielding. Ecology 91, 2213–2220.
Hillebrand, H., Bennett, D.M., Cadotte, M.W., 2008. Consequences of dominance: a

28
C. Wang, et al. Acta Oecologica 98 (2019) 25–29

review of evenness effects on local and regional ecosystem processes. Ecology 89, Simpson, E.H., 1949. Measurement of diversity. Nature 163, 688.
1510–1520. Tilman, D., Downing, J.A., 1994. Biodiversity and stability in grasslands. Nature 367,
Hu, Z.Z., 2000. Grass Industry Development and Ecological Environment in Tibet Plateau. 363–365.
China Tibetology Press, Beijing, pp. 1–124. Tilman, D., Isbell, F., Cowles, J.M., 2014. Biodiversity and ecosystem functioning. Annu.
Isbell, F.I., Calcagno, V., Hector, A., Connolly, J., Harpole, W.S., Reich, P.B., Rev. Ecol. Evol. Syst. 45, 471–493.
Scherer‒Lorenzen, M., Schmid, B., Tilman, D., van Ruijven, J., Weigelt, A., Wilsey, Valone, T.J., Balaban Feld, J., 2018. Impact of exotic invasion on the temporal stability of
B.J., Zavaleta, E.S., Loreau, M., 2011. High plant diversity is needed to maintain natural annual plant communities. Oikos 127, 56–62.
ecosystem services. Nature 477, 199–202. Wang, C.Y., Jiang, K., Liu, J., Zhou, J.W., Wu, B.D., 2018a. Moderate and heavy Solidago
Isbell, F.I., Polley, H.W., Wilsey, B.J., 2009. Biodiversity, productivity and the temporal canadensis L. invasion are associated with decreased taxonomic diversity but in-
stability of productivity: patterns and processes. Ecol. Lett. 12, 443–451. creased functional diversity of plant communities in East China. Ecol. Eng. 112,
Ives, A.R., Carpenter, S.R., 2007. Stability and diversity of ecosystems. Science 317, 55–64.
58–62. Wang, C.Y., Jiang, K., Zhou, J.W., Wu, B.D., 2018b. Solidago canadensis invasion affects
Lehman, C.L., Tilman, D., 2000. Biodiversity, stability, and productivity in competitive soil N‒fixing bacterial communities in heterogeneous landscapes in urban ecosystems
communities. Am. Nat. 156, 534–552. in East China. Sci. Total Environ. 631‒632, 702–713.
Loreau, M., De Mazancourt, C., 2013. Biodiversity and ecosystem stability: a synthesis of Wang, D.J., 2015. Tibet Yearbook. Written by Editorial Board of Tibet Yearbook. Tibet
underlying mechanisms. Ecol. Lett. 16, 106–115. People’s Publishing House, Lasa, pp. 455.
Margalef, R., 1951. Diversidad de especies en las comunidades naturales, vol. 6. Wen, L., Dong, S.K., Zhu, L., Li, X.Y., Shi, J.J., Wang, Y.L., Ma, Y.S., 2010. The con-
Publicaciones del Instituto de Biologia Aplicada, Barcelona, pp. 59–72. struction of grassland degradation index for alpine meadow in qinghai‒Tibetan
Melese, S.M., Ayele, B., 2017. Woody plant diversity, structure and regeneration in the plateau. Proc. Environ. Sci. 2, 1966–1969.
ambo state forest, south gondar zone, northwest Ethiopia. J. For. Res. 28, 133–144. Yang, H.J., Jiang, L., Li, L.H., Li, A., Wu, M.Y., Wan, S.Q., 2012. Diversity‒dependent
Pielou, E.C., 1966. The measurement of diversity in different types of biological collec- stability under mowing and nutrient addition: evidence from a 7‒year grassland
tions. J. Theor. Biol. 13, 131–144. experiment. Ecol. Lett. 15, 619–626.
Reeder, J.D., Schuman, G.E., 2002. Influence of livestock grazing on C sequestration in Yang, Z.L., van Ruijven, J., Du, G.Z., 2011. The effects of long‒term fertilization on the
semi‒arid mixed‒grass and short‒grass rangelands. Environ. Pollut. 116, 457–463. temporal stability of alpine meadow communities. Plant Soil 345, 315–324.
Rong, X.S., He, M., Wang, C.Y., Li, Y., Dai, Z.C., Liang, Z.S., Wan, L.Y., Gao, Z.Y., Jiang, K., Yang, Z.L., Zhang, Q., Su, F.L., Zhang, C.H., Pu, Z.C., Xia, J.Y., Wan, S.Q., Jiang, L., 2017.
Wu, B.D., Zhang, L., Zhou, J.W., 2018. Analysis on the diversity of soil bacterial and Daytime warming lowers community temporal stability by reducing the abundance of
fungal communities in the degraded Alpine Grassland in Northern Tibet Plateau. dominant, stable species. Glob. Chang. Biol. 23, 154–163.
Ecol. Environ. Sci. 27, 1646–1651 (In Chinese). Zavaleta, E.S., Pasari, J.R., Hulvey, K.B., Tilman, G.D., 2010. Sustaining multiple eco-
Sasaki, T., Lauenroth, W.K., 2011. Dominant species, rather than diversity, regulates system functions in grassland communities requires higher biodiversity. Proc. Natl.
temporal stability of plant communities. Oecologia 166, 761–768. Acad. Sci. U.S.A. 107, 1443–1446.
Shannon, C.E., Weaver, W., 1949. The Mathematical Theory of Communication. Zhao, G.F., Yu, C.Q., Wu, J.X., Luo, L.M., Miao, Y.J., 2013. Research progress on re-
University of Illinois Press, Urbana, Illinois, pp. 1–117. storation and management of degraded alpine meadow in Qinahai‒Tibet Plateau.
Shao, W., Cai, X.B., 2008. Grassland degradation and its formation causes analysis in Guizhou Agr. Sci. 41, 125–129 (In Chinese).
Tibetan Plateau. Int. Sci. Soil Water Conserv. 6, 112–116 (In Chinese).

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