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Journal of Arid Land  2019, Vol. 11 Issue (4): 595-607    DOI: 10.1007/s40333-019-0122-6
Community phylogenetic structure of grasslandsand its relationship with environmental factors on the Mongolian Plateau
Lei DONG1, Cunzhu LIANG1,*(), Frank Yonghong LI1, Liqing ZHAO1, Wenhong MA1, Lixin WANG1, Lu WEN1, Ying ZHENG1, Zijing LI1, Chenguang ZHAO2, IndreeTUVSHINTOGTOKH3
1Ministry of Education Key Laboratory of Ecology and Resource Use of the Mongolian Plateau & Inner Mongolia Key Laboratory of Grassland Ecology, School of Ecology and Environment, Inner Mongolia University, Hohhot 010021, China;
2 Forestry and DesertControl Research Institute of Alagxa League,Bayanhot750306,China
3Institute of General and Experimental Biology, Mongolian Academy of Sciences, Ulaanbaatar-51, Mongolia
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The community assembly rules and species coexistence have always been interested by ecologists.The community phylogenetic structure is the consequence of the interaction process between the organisms and the abiotic environmentand has been used to explain the relative impactof abiotic and biotic factors on species co-existence. In recent years, grassland degradation and biodiversity loss have become increasingly severe on the Mongolian Plateau,while the drivers for these changes are not clearly explored, especially whether climate change is a main factor is debated in academia. In this study, we examined the phylogenetic structure of grassland communities along five transects of climate aridity on the Mongolian Plateau, and analyzed their relations with environmental factors, with the aims to understand theformation mechanismof the grasslandcommunities and the role of climatic factors.We surveyed grassland communities at 81 sites along the five transects,and calculated theirnet relatedness index (NRI)attwo different quadratscales (smallscale of 1 m2 and largescale of 5 m2) to characterize thecommunity phylogenetic structure andanalyzeitsrelationship withthe key 11 environmental factors. We also calculatedthe generalized UniFrac distance (GUniFrac)among the grassland communities to quantify the influence of spatial distance and environmental distance on the phylogenetic β diversity. The results indicated that plant community survey using the largescalequadrat contained sufficient species to represent community compositions. The community phylogenetic structure of grasslandswas significantly overdispersed at both the small and largescales, and the degree of overdispersion was greater at the large scalethan at the smallscale, suggesting that competitive exclusion instead of habitat filtering played a major role in determination of community composition. Altitude was the main factor affecting the community phylogenetic structure, whereas climatic factors, such as precipitation and temperature, had limited influence. Theprincipal component analysis of the 11 environmental factorsrevealed that 94.04% of their variation was accounted by the first four principal components.Moreover only 14.29% and 23.26% of the variation in community phylogenetic structurewere explained by the first four principal componentsat the small and largescales, respectively. Phylogenetic β diversity was slightly significantly correlated with both spatial distance and environmental distance, however,environmental distancehada less explanatory powerthan spatial distance, indicating a limited environmental effect on the community phylogenetic structure of grasslands on the Mongolian Plateau. In view of the limited effect of climatic factors on the community phylogenetic structure of grasslands, climate change may have a smaller impact on grassland degradation than previously thought.

Key wordsphylogenetic overdispersion      environmental factors      phylogenetic β diversity;      spatial scale      environmental distance      climate change      Mongolian Plateau     
Received: 30 March 2018      Published: 10 August 2019
Fund:  This study was supported by the National Key Research and Development Program of China (2016YFC0500503) and the Science and Technology Program of Inner Mongolia Autonomous Region of China (20140409, 201503001). The authors also thank to the reviewers and editors for their helpful suggestions.
Corresponding Authors: Cunzhu LIANG     E-mail:
About author:

The first and second authors contributed equally to this work.

Cite this article:

Lei DONG, Cunzhu LIANG, Frank Yonghong LI, Liqing ZHAO, Wenhong MA, Lixin WANG, Lu WEN, Ying ZHENG, Zijing LI, Chenguang ZHAO, IndreeTUVSHINTOGTOKH. Community phylogenetic structure of grasslandsand its relationship with environmental factors on the Mongolian Plateau. Journal of Arid Land, 2019, 11(4): 595-607.

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[1] Bai Y, Han X, Wu J, et al.2004. Ecosystem stability and compensatory effects in the Inner Mongolia grassland. Nature, 431(7005):181-184.
[2] Bai Y, Wu J, Xing Q, et al.2008. Primary production and rain use efficiency across a precipitation gradient on the Mongolia Plateau. Ecology, 89(8):2140-2153.
[3] Bao G, Qin Z, Bao Y, et al.2014. NDVI-Based long-term vegetation dynamics and its response to climatic change in the Mongolian Plateau. Remote Sensing, 6(9):8337-8358.
[4] Baraloto C, Hardy OJ, Paine C E T, et al.2012. Using functional traits and phylogenetic trees to examine the assembly of tropical tree communities. Journal of Ecology, 100(3):690-701.
[5] Buckley LB, Kingsolver JG.2012. Functional and phylogenetic approaches to forecasting species' responses to climate change. Annual Review of Ecology, Evolution, and Systematics, 43(1):205-226.
[6] Burns JH, Strauss SY.2011. More closely related species are more ecologically similar in an experimental test. Proceedings of the National Academy of Sciences of the United States of America, 108(13):5302-5307.
[7] Cadotte MW, Dinnage R, Tilman D.2012. Phylogenetic diversity promotes ecosystem stability. Ecology, 93(Suppl.8):S223-S233.
[8] Cao H, Zhao X, Wang S, et al.2015. Grazing intensifies degradation of a Tibetan Plateau alpine meadow through plant-pest interaction. Ecology and Evolution, 5(12):2478-2486.
[9] Cavender-Bares J, Ackerly D D, Baum D A, et al.2004. Phylogenetic overdispersion in Floridian oak communities. The American Naturalist, 163(6):823-843.
[10] Cavender-Bares J, Keen A, Miles B.2006. Phylogenetic structure of Floridian plant communities depends on taxonomic and spatial scale. Ecology, 87(Suppl.7):S109-S122.
[11] Cavender-Bares J, Kozak KH, Fine PV, et al.2009. The merging of community ecology and phylogenetic biology. Ecology Letters, 12(7):693-715.
[12] Chen B, Zhang X, Tao J, et al.2014. The impact of climate change and anthropogenic activities on alpine grassland over the Qinghai-Tibet Plateau. Agricultural and Forest Meteorology, 189-190:11-18.
[13] Chen J, Huang D, Shiyomi M, et al.2007. Spatial heterogeneity and diversity of vegetation at the landscape level in Inner Mongolia, China, with special reference to water resources. Landscape and Urban Planning, 82(4):222-232.
[14] Chen J, Bittinger K, Charlson ES, et al.2012. Associating microbiome composition with environmental covariates using generalized UniFrac distances. Bioinformatics, 28(16):2106-2113.
[15] Chu C, Bartlett M, Wang Y, et al.2016. Does climate directly influence NPP globally? Global Change Biology, 22(1):12-24.
[16] Cramer W, Bondeau A, Woodward FI, et al.2001. Global response of terrestrial ecosystem structure and function to CO2 and climate change: results from six dynamic global vegetation models. Global Change Biology, 7(4):357-373.
[17] Critchley C N R, Adamson HF, McLean B M L, et al.2008. Vegetation dynamics and livestock performance in system-scale studies of sheep and cattle grazing on degraded upland wet heath. Agriculture, Ecosystems & Environment, 128(1-2):59-67.
[18] Davies KF, Chesson P, Harrison S, et al.2005. Spatial heterogeneity explains the scale dependence of the native-exotic diversity relationship. Ecology, 86(6):1602-1610.
[19] De Mendiburu F.2014. Agricolae: statistical procedures for agricultural research. R package version1.3-0. Lima: National Engineering University.
[20] Donoghue MJ.2008. A phylogenetic perspective on the distribution of plant diversity. Proceedings of the National Academy of Sciences, 105(Suppl. 1):11549-11555.
[21] Du M, Kawashima S, Yonemura S, et al.2004. Mutual influence between human activities and climate change in the Tibetan Plateau during recent years. Global and Planetary Change, 41(3-4):241-249.
[22] Emerson BC, Gillespie RG.2008. Phylogenetic analysis of community assembly and structure over space and time. Trends in Ecology &Evolution, 23(11):619-630.
[23] Fang J, Wang X, Shen Z, et al.2009. Methods and protocols for plant community inventory. Biodiversity Science, 17(6):533-548.
[24] Feng G, Mi X, Eiserhardt WL, et al.2015. Assembly of forest communities across East Asia-insights from phylogenetic community structure and species pool scaling. Scientific Reports, 5:9337, doi: org/9310.1038/srep09337.
[25] Fernández RJ.2002. Do humans create deserts? Trends in Ecology & Evolution, 17(1):6-7.
[26] Field CB, Behrenfeld MJ, Randerson JT, et al.1998. Primary production of the biosphere: integrating terrestrial and oceanic components. Science, 281(5374):237-240.
[27] Freestone AL, Inouye BD.2006. Dispersal limitation and environmental heterogeneity shape scale-dependent diversity patterns in plant communities. Ecology, 87(10):2425-2432.
[28] Gang C, Zhou W, Chen Y, et al.2014. Quantitative assessment of the contributions of climate change and human activities on global grassland degradation. Environmental Earth Sciences, 72(11):4273-4282.
[29] Gerhold P, Pärtel M, Liira J, et al.2008. Phylogenetic structure of local communities predicts the size of the regional species pool. Journal of Ecology, 96(4):709-712.
[30] Hijmans RJ, Cameron SE, Parra JL, et al.2005. Very high resolution interpolated climate surfaces for global land areas. International Journal of Climatology, 25(15):1965-1978.
[31] Hilker T, Natsagdorj E, Waring RH, et al.2014. Satellite observed widespread decline in Mongolian grasslands largely due to overgrazing. Global Change Biology, 20(2):418-428.
[32] Hoiss B, Krauss J, Potts SG, et al.2012. Altitude acts as an environmental filter on phylogenetic composition, traits and diversity in bee communities. Proceedings of the Royal Society B: Biological Sciences, 279(1746):4447-4456.
[33] Hughes JW, Fahey TJ, Bormann FH.1988. Population persistence and reproductive ecology of a forest herb: Aster acuminatus. American Journal of Botany, 75(7):1057-1064.
[34] Kembel SW, Hubbell SP.2006. The phylogenetic structure of a neotropical forest tree community. Ecology, 87(Suppl. 7):86-99.
[35] Kembel SW, Cowan PD, Helmus MR, et al.2010. Picante: R tools for integrating phylogenies and ecology. Bioinformatics, 26(11):1463-1466.
[36] Kraft NJ B, Cornwell WK, Webb CO, et al.2007. Trait evolution, community assembly, and the phylogenetic structure of ecological communities. The American Naturalist, 170(2):271-283.
[37] Kraft NJB, Ackerly DD.2010. Functional trait and phylogenetic tests of community assembly across spatial scales in an Amazonian forest. Ecological Monographs, 80(3):401-422.
[38] Kraft NJ B, Comita LS, Chase JM, et al.2011. Disentangling the drivers of β diversity along latitudinal and elevational gradients. Science, 333(6050):1755-1758.
[39] Lavergne S, Mouquet N, Thuiller W, et al.2010. Biodiversity and climate change: integrating evolutionary and ecological responses of species and communities. Annual Review of Ecology, Evolution, and Systematics, 41(1):321-350.
[40] Letcher SG.2010. Phylogenetic structure of angiosperm communities during tropical forest succession. Proceedings of the Royal Society B: Biological Sciences, 277(1678):97-104.
[41] Lin G, Huang Z, Lin Z, et al.2010. Beta diversity of forest community on Dinghushan. Acta Ecologica Sinica, 30(18):4875-4880. (in Chinese)
[42] Lu Y, Zhuang Q, Zhou G, et al.2009. Possible decline of the carbon sink in the Mongolian Plateau during the 21st century. Environmental Research Letters, 4(4):045023.
[43] McKinney ML.2002. Urbanization, biodiversity, and conservation: the impacts of urbanization on native species are poorly studied, but educating a highly urbanized human population about these impacts can greatly improve species conservation in all ecosystems. BioScience, 52(10):883-890.
[44] Melillo JM, McGuire AD, Kicklighter DW, et al.1993. Global climate change and terrestrial net primary production. Nature, 363(6426):234-240.
[45] Nishitani S, Takada T, Kachi N.1999. Optimal resource allocation to seeds and vegetative propagules under density-dependent regulation in Syneilesis palmata (Compositae). Plant Ecology, 141(1-2):179-189.
[46] Qian H, Hao Z, Zhang J.2014. Phylogenetic structure and phylogenetic diversity of angiosperm assemblages in forests along an elevational gradient in Changbaishan, China. Journal of Plant Ecology, 7(2):154-165.
[47] Qian H, Jin Y, Ricklefs RE.2017. Phylogenetic diversity anomaly in angiosperms between eastern Asia and eastern North America. Proceedings of the National Academy of Sciences, 114(43):11452-11457.
[48] Ricklefs RE, Latham RE.1992. Intercontinental correlation of geographical ranges suggests stasis in ecological traits of relict genera of temperate perennial herbs. The American Naturalist, 139(6):1305-1321.
[49] Roughgarden J.1983. Competition and theory in community ecology. The American Naturalist, 122(5):583-601.
[50] Slingsby JA, Verboom GA.2006. Phylogenetic relatedness limits co-occurrence at fine spatial scales: evidence from the schoenoid sedges (Cyperaceae: Schoeneae) of the cape floristic region, South Africa. The American Naturalist, 168(1):14-27.
[51] Soliveres S, Torices R, Maestre FT.2012. Environmental conditions and biotic interactions acting together promote phylogenetic randomness in semi-arid plant communities: new methods help to avoid misleading conclusions. Journal of Vegetation Science, 23(5):822-836.
[52] Stohlgren TJ, Falkner M, Schell L.1995. A modified-Whittaker nested vegetation sampling method. Vegetatio, 117(2):113-121.
[53] Swenson NG, Enquist BJ, Pither J, et al.2006. The problem and promise of scale dependency in community phylogenetics. Ecology, 87(10):2418-2424.
[54] Swenson NG, Enquist BJ, Thompson J, et al.2007. The influence of spatial and size scale on phylogenetic relatedness in tropical forest communities. Ecology, 88(7):1770-1780.
[55] Tamm CO.1956. Further observations on the survival and flowering of some perennial herbs, I. Oikos, 7(2):273-292.
[56] The Angiosperm Phylogeny Group.2009. An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG III. Botanical Journal of the Linnean Society, 161(2):105-121.
[57] Tilman D.2004. Niche tradeoffs, neutrality, and community structure: a stochastic theory of resource competition, invasion, and community assembly. Proceedings of the National Academy of Sciences of the United States of America, 101(30):10854-10861.
[58] Vamosi S M, Heard SB, Vamosi JC, et al.2009. Emerging patterns in the comparative analysis of phylogenetic community structure. Molecular Ecology, 18(4):572-592.
[59] Vandandorj S, Gantsetseg B, Boldgiv B.2015. Spatial and temporal variability in vegetation cover of Mongolia and its implications. Journal of Arid Land, 7(4):450-461.
[60] Volkov I, Banavar JR, He F, et al.2005. Density dependence explains tree species abundance and diversity in tropical forests. Nature, 438(7068):658-661.
[61] Webb CO.2000. Exploring the phylogenetic structure of ecological communities: an example for rain forest trees. The American Naturalist, 156(2):145-155.
[62] Webb CO, Ackerly DD, Mcpeek MA, et al.2002. Phylogenies and community ecology. Annual Review of Ecology and Systematics, 33(1):475-505.
[63] Webb CO, Donoghue MJ.2005. Phylomatic: tree assembly for applied phylogenetics. Molecular Ecology Notes, 5(1):181-183.
[64] Webb CO, Gilbert GS, Donoghue MJ.2006. Phylodiversity-dependent seedling mortality, size structure, and disease in a Bornean rain forest. Ecology, 87(Suppl. 7):S123-S131.
[65] Williams NSG, Morgan JW, Mcdonnell MJ, et al.2005. Plant traits and local extinctions in natural grasslands along an urban-rural gradient. Journal of Ecology, 93(6):1203-1213.
[66] Willis CG, Ruhfel B, Primack RB, et al.2008. Phylogenetic patterns of species loss in Thoreau's woods are driven by climate change. Proceedings of the National Academy of Sciences, 105(44):17029-17033.
[67] Willis CG, Halina M, Lehman C, et al.2010. Phylogenetic community structure in Minnesota oak savanna is influenced by spatial extent and environmental variation. Ecography, 33(3):565-577.
[68] Yang H, Wu M, Liu W, et al.2011. Community structure and composition in response to climate change in a temperate steppe. Global Change Biology, 17(1):452-465.
[69] Yang X, Yang Z, Tan J, et al.2018. Nitrogen fertilization, not water addition, alters plant phylogenetic community structure in a semi-arid steppe. Journal of Ecology, 106(3):991-1000.
[70] Yue X.2011. Study on the flora of seed plants in the Mongolian Plateau.MSc Thesis. Hohhot: Inner Mongolia Agricultural University. (in Chinese)
[71] Zavaleta ES, Shaw MR, Chiariello NR, et al.2003. Additive effects of simulated climate changes, elevated CO2, and nitrogen deposition on grassland diversity. Proceedings of the National academy of Sciences, 100(13):7650-7654.
[72] Zhang G, Kang Y, Han G, et al.2011. Effect of climate change over the past half century on the distribution, extent and NPP of ecosystems of Inner Mongolia. Global Change Biology, 17(1):377-389.
[73] Zhang P P, Shao M A, Zhang X C.2017. Spatial pattern of plant species diversity and the influencing factors in a Gobi Desert within the Heihe River Basin, Northwest China. Journal of Arid Land, 9(3): 379-393.
[74] Zhao X, Hu H, Shen H, et al.2015. Satellite-indicated long-term vegetation changes and their drivers on the Mongolian Plateau. Landscape Eology, 30(9):1599-1611.
[75] Zhou X, Yamaguchi Y, Arjasakusuma S.2017. Distinguishing the vegetation dynamics induced by anthropogenic factors using vegetation optical depth and AVHRR NDVI: A cross-border study on the Mongolian Plateau. Science of the Total Environment, 616-617:730-743.
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