Community phylogenetic structure of grasslands and its relationship with environmental factors on the Mongolian Plateau

  • Lei Dong
  • Cunzhu LiangEmail author
  • Frank Yonghong Li
  • Liqing Zhao
  • Wenhong Ma
  • Lixin Wang
  • Lu Wen
  • Ying Zheng
  • Zijing Li
  • Chenguang Zhao
  • Indree Tuvshintogtokh


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 environment and has been used to explain the relative impact of 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 the formation mechanism of the grassland communities and the role of climatic factors. We surveyed grassland communities at 81 sites along the five transects, and calculated their net relatedness index (NRI) at two different quadrat scales (small scale of 1 m2 and large scale of 5 m2) to characterize the community phylogenetic structure and analyze its relationship with the key 11 environmental factors. We also calculated the 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 large scale quadrat contained sufficient species to represent community compositions. The community phylogenetic structure of grasslands was significantly overdispersed at both the small and large scales, and the degree of overdispersion was greater at the large scale than at the small scale, 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. The principal component analysis of the 11 environmental factors revealed 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 structure were explained by the first four principal components at the small and large scales, respectively. Phylogenetic β diversity was slightly significantly correlated with both spatial distance and environmental distance, however, environmental distance had a less explanatory power than 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.


phylogenetic overdispersion environmental factors phylogenetic β diversity spatial scale environmental distance climate change Mongolian Plateau 


Unable to display preview. Download preview PDF.

Unable to display preview. Download preview PDF.


  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.Google Scholar
  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.Google Scholar
  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.Google Scholar
  4. Baraloto C, Hardy O J, 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.Google Scholar
  5. Buckley L B, Kingsolver J G. 2012. Functional and phylogenetic approaches to forecasting species’ responses to climate change. Annual Review of Ecology, Evolution, and Systematics, 43(1): 205–226.Google Scholar
  6. Burns J H, Strauss S Y. 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.Google Scholar
  7. Cadotte M W, Dinnage R, Tilman D. 2012. Phylogenetic diversity promotes ecosystem stability. Ecology, 93(Suppl. 8): S223–S233.Google Scholar
  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.Google Scholar
  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.Google Scholar
  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.Google Scholar
  11. Cavender-Bares J, Kozak K H, Fine P V, et al. 2009. The merging of community ecology and phylogenetic biology. Ecology Letters, 12(7): 693–715.Google Scholar
  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.Google Scholar
  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.Google Scholar
  14. Chen J, Bittinger K, Charlson E S, et al. 2012. Associating microbiome composition with environmental covariates using generalized UniFrac distances. Bioinformatics, 28(16): 2106–2113.Google Scholar
  15. Chu C, Bartlett M, Wang Y, et al. 2016. Does climate directly influence NPP globally? Global Change Biology, 22(1): 12–24.Google Scholar
  16. Cramer W, Bondeau A, Woodward F I, 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.Google Scholar
  17. Critchley C N R, Adamson H F, 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.Google Scholar
  18. Davies K F, Chesson P, Harrison S, et al. 2005. Spatial heterogeneity explains the scale dependence of the native-exotic diversity relationship. Ecology, 86(6): 1602–1610.Google Scholar
  19. De Mendiburu F. 2014. Agricolae: statistical procedures for agricultural research. R package version 1.3-0. Lima: National Engineering University.Google Scholar
  20. Donoghue M J. 2008. A phylogenetic perspective on the distribution of plant diversity. Proceedings of the National Academy of Sciences, 105 (Suppl. 1): 11549–11555.Google Scholar
  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.Google Scholar
  22. Emerson B C, Gillespie R G. 2008. Phylogenetic analysis of community assembly and structure over space and time. Trends in Ecology & Evolution, 23(11): 619–630.Google Scholar
  23. Fang J, Wang X, Shen Z, et al. 2009. Methods and protocols for plant community inventory. Biodiversity Science, 17(6): 533–548.Google Scholar
  24. Feng G, Mi X, Eiserhardt W L, 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.Google Scholar
  25. Fernandez R J. 2002. Do humans create deserts? Trends in Ecology & Evolution, 17(1): 6–7.Google Scholar
  26. Field C B, Behrenfeld M J, Randerson J T, et al. 1998. Primary production of the biosphere: integrating terrestrial and oceanic components. Science, 281(5374): 237–240.Google Scholar
  27. Freestone A L, Inouye B D. 2006. Dispersal limitation and environmental heterogeneity shape scale-dependent diversity patterns in plant communities. Ecology, 87(10): 2425–2432.Google Scholar
  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.Google Scholar
  29. Gerhold P, Partel 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.Google Scholar
  30. Hijmans R J, Cameron S E, Parra J L, et al. 2005. Very high resolution interpolated climate surfaces for global land areas. International Journal of Climatology, 25(15): 1965–1978.Google Scholar
  31. Hilker T, Natsagdorj E, Waring R H, et al. 2014. Satellite observed widespread decline in Mongolian grasslands largely due to overgrazing. Global Change Biology, 20(2): 418–428.Google Scholar
  32. Hoiss B, Krauss J, Potts S G, 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.Google Scholar
  33. Hughes J W, Fahey T J, Bormann F H. 1988. Population persistence and reproductive ecology of a forest herb: Aster acuminatus. American Journal of Botany, 75(7): 1057–1064.Google Scholar
  34. Kembel S W, Hubbell S P. 2006. The phylogenetic structure of a neotropical forest tree community. Ecology, 87(Suppl. 7): 86–99.Google Scholar
  35. Kembel S W, Cowan P D, Helmus M R, et al. 2010. Picante: R tools for integrating phylogenies and ecology. Bioinformatics, 26(11): 1463–1466.Google Scholar
  36. Kraft N J B, Cornwell W K, Webb C O, et al. 2007. Trait evolution, community assembly, and the phylogenetic structure of ecological communities. The American Naturalist, 170(2): 271–283.Google Scholar
  37. Kraft N J B, Ackerly D D. 2010. Functional trait and phylogenetic tests of community assembly across spatial scales in an Amazonian forest. Ecological Monographs, 80(3): 401–422.Google Scholar
  38. Kraft N J B, Comita L S, Chase J M, et al. 2011. Disentangling the drivers of β diversity along latitudinal and elevational gradients. Science, 333(6050): 1755–1758.Google Scholar
  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.Google Scholar
  40. Letcher S G. 2010. Phylogenetic structure of angiosperm communities during tropical forest succession. Proceedings of the Royal Society B: Biological Sciences, 277(1678): 97–104.Google Scholar
  41. Lin G, Huang Z, Lin Z, et al. 2010. Beta diversity of forest community on Dinghushan. Acta Ecologica Sinica, 30(18): 4875–4880.Google Scholar
  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.Google Scholar
  43. McKinney M L. 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.Google Scholar
  44. Melillo J M, McGuire A D, Kicklighter D W, et al. 1993. Global climate change and terrestrial net primary production. Nature, 363(6426): 234–240.Google Scholar
  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.Google Scholar
  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.Google Scholar
  47. Qian H, Jin Y, Ricklefs R E. 2017. Phylogenetic diversity anomaly in angiosperms between eastern Asia and eastern North America. Proceedings of the National Academy of Sciences, 114(43): 11452–11457.Google Scholar
  48. Ricklefs R E, Latham R E. 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.Google Scholar
  49. Roughgarden J. 1983. Competition and theory in community ecology. The American Naturalist, 122(5): 583–601.Google Scholar
  50. Slingsby J A, Verboom G A. 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.Google Scholar
  51. Soliveres S, Torices R, Maestre F T. 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.Google Scholar
  52. Stohlgren T J, Falkner M, Schell L. 1995. A modified-Whittaker nested vegetation sampling method. Vegetatio, 117(2): 113–121.Google Scholar
  53. Swenson N G, Enquist B J, Pither J, et al. 2006. The problem and promise of scale dependency in community phylogenetics. Ecology, 87(10): 2418–2424.Google Scholar
  54. Swenson N G, Enquist B J, Thompson J, et al. 2007. The influence of spatial and size scale on phylogenetic relatedness in tropical forest communities. Ecology, 88(7): 1770–1780.Google Scholar
  55. Tamm C O. 1956. Further observations on the survival and flowering of some perennial herbs, I. Oikos, 7(2): 273–292.Google Scholar
  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.Google Scholar
  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.Google Scholar
  58. Vamosi S M, Heard S B, Vamosi J C, et al. 2009. Emerging patterns in the comparative analysis of phylogenetic community structure. Molecular Ecology, 18(4): 572–592.Google Scholar
  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.Google Scholar
  60. Volkov I, Banavar J R, He F, et al. 2005. Density dependence explains tree species abundance and diversity in tropical forests. Nature, 438(7068): 658–661.Google Scholar
  61. Webb C O. 2000. Exploring the phylogenetic structure of ecological communities: an example for rain forest trees. The American Naturalist, 156(2): 145–155.Google Scholar
  62. Webb C O, Ackerly D D, Mcpeek M A, et al. 2002. Phylogenies and community ecology. Annual Review of Ecology and Systematics, 33(1): 475–505.Google Scholar
  63. Webb C O, Donoghue M J. 2005. Phylomatic: tree assembly for applied phylogenetics. Molecular Ecology Notes, 5(1): 181–183.Google Scholar
  64. Webb C O, Gilbert G S, Donoghue M J. 2006. Phylodiversity-dependent seedling mortality, size structure, and disease in a Bornean rain forest. Ecology, 87(Suppl. 7): S123–S131.Google Scholar
  65. Williams N S G, Morgan J W, Mcdonnell M J, et al. 2005. Plant traits and local extinctions in natural grasslands along an urban-rural gradient. Journal of Ecology, 93(6): 1203–1213.Google Scholar
  66. Willis C G, Ruhfel B, Primack R B, 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.Google Scholar
  67. Willis C G, 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.Google Scholar
  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.Google Scholar
  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.Google Scholar
  70. Yue X. 2011. Study on the flora of seed plants in the Mongolian Plateau. MSc Thesis. Hohhot: Inner Mongolia Agricultural University. (in Chinese)Google Scholar
  71. Zavaleta E S, Shaw M R, Chiariello N R, 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.Google Scholar
  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.Google Scholar
  73. 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.Google Scholar
  74. 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.Google Scholar

Copyright information

© Xinjiang Institute of Ecology and Geography, Chinese Academy of Sciences, Science Press and Springer-Verlag GmbH Germany, part of Springer Nature 2019

Authors and Affiliations

  • Lei Dong
    • 1
  • Cunzhu Liang
    • 1
    Email author
  • Frank Yonghong Li
    • 1
  • Liqing Zhao
    • 1
  • Wenhong Ma
    • 1
  • Lixin Wang
    • 1
  • Lu Wen
    • 1
  • Ying Zheng
    • 1
  • Zijing Li
    • 1
  • Chenguang Zhao
    • 2
  • Indree Tuvshintogtokh
    • 3
  1. 1.Ministry of Education Key Laboratory of Ecology and Resource Use of the Mongolian Plateau & Inner Mongolia Key Laboratory of Grassland Ecology, School of Ecology and EnvironmentInner Mongolia UniversityHohhotChina
  2. 2.Forestry and Desert Control Research Institute of Alagxa LeagueBayanhotChina
  3. 3.Institute of General and Experimental BiologyMongolian Academy of SciencesUlaanbaatar-51Mongolia

Personalised recommendations