Community Ecology

, Volume 16, Issue 2, pp 215–222 | Cite as

Interplay of temperature and woody cover shapes herb communities along an elevational gradient in a temperate forest in Beijing, China

  • Zihan Jiang
  • Keming MaEmail author
  • M. Anand
  • Yuxin Zhang


Abiotic and biotic factors have the potential to alter herb communities, however, few studies consider feedback between them. This study explores how variation of species interaction and climatic conditions associated with changes in altitude affect herb community composition. We sampled accumulated temperatures of growth duration (June-November) (ATGD), maximum summer temperatures (MST) and herb community composition (herb height, abundance, richness) on non-gaps and forest-gaps site across an elevational gradient. A significant negative relationship was detected between MST and herb richness. The temperature of non-gaps was cooler than that of forest gaps, and overstory cover positively correlated with herb abundance. However, the ATGD exhibited a negative relationship with overstory cover, in that overstory cover decreased with ATGD. We suggested that temperature has a profound effect on height and richness of herb communities, while the overstory cover is moderating the effect of temperature on herb community structure and influence the abundance of herb community. Conversely, decreases in ATGD weakened the relative importance of overstory cover. We concluded that the interaction of temperature and overstory cover shapes the morphology, abundance and richness of herb communities.


Accumulated temperature of growth duration Elevational gradient Herb communities Maximum summer temperature Overstory cover 



Accumulated Temperatures of Growth Duration


Maximum Summer Temperatures


Overstory Cover


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  1. Ashton, P.S. 2003. Floristic zonation of tree communities on wet tropical mountains revisited. Perspect. Plant Ecol. 6: 87–104.CrossRefGoogle Scholar
  2. Bao, S. 2000. Analysis Method of Soil Agricultural Chemistry. China Agr. Press, Beijing.Google Scholar
  3. Bartels, S.F. and H.Y.H. Chen. 2013. Interactions between overstory and understory vegetation along an overstory compositional gradient. J. Veg. Sci. 24: 543–552.CrossRefGoogle Scholar
  4. Bauhus, J. and N. Bartsch. 1996. Fine-root growth in beech (Fagus sylvatica) forest gaps. Can. J. Forest. Res. 26: 2153–2159.CrossRefGoogle Scholar
  5. Boggs, C.L. and D.D. Murphy. 1997. Community composition in mountain ecosystems: climatic determinants of montane butterfly distributions. Global Ecol. Biogeogr. 6: 39–48.CrossRefGoogle Scholar
  6. Bowles, M.L., K.A. Jacobs and J.L. Mengler. 2007. Long-term changes in an oak forest’s woody understory and herb layer with repeated burning. J. Torrey Bot. Soc. 134: 223–237.CrossRefGoogle Scholar
  7. Burton, J.I., D.J. Mladenoff, J.A. Forrester, and M.K. Clayton. 2014. Experimentally linking disturbance, resources and productivity to diversity in forest ground-layer plant communities. J. Ecol. 102: 1634–1648.CrossRefGoogle Scholar
  8. Caldeira, M.C., I. Ibanez, C. Nogueira, M.N. Bugalho, X. Lecomte, A. Moreira and J.S. Pereira. 2014. Direct and indirect effects of tree canopy facilitation in the recruitment of Mediterranean oaks. J. Appl. Ecol. 51: 349–358.CrossRefGoogle Scholar
  9. Cairns, M.A., S. Brown, E.H. Helmer and G.A. Baumgardner. 1997. Root biomass allocation in the world’s upland forests. Oecologia 111: 1–11.CrossRefPubMedPubMedCentralGoogle Scholar
  10. Colwell, R.K., G. Brehm, C.L. Cardelus, A.C. Gilman and J.T. Longino. 2008. Global warming, elevational range shifts, and lowland biotic attrition in the wet tropics. Science 322: 258–261.CrossRefPubMedPubMedCentralGoogle Scholar
  11. Crozier, C.R. and R.E. Boerner. 1984. Correlations of understory herb distribution patterns with microhabitats under different tree species in a mixed mesophytic forest. Oecologia 62: 337–343.CrossRefPubMedPubMedCentralGoogle Scholar
  12. Daubenmire, R. 1943. Soil temperature versus drought as a factor determining lower altitudinal limits of trees in the Rocky Mountains. Bot. Gaz. 105: 1–13.CrossRefGoogle Scholar
  13. De Frenne, P. et al. 2013. Microclimate moderates plant responses to macroclimate warming. P. Natl. Acad. of. Sci. USA. 110: 18561– 18565.CrossRefGoogle Scholar
  14. Denno, R.F., G.K. Roderick, M.A. Peterson, A.F. Huberty, H.G. Dobel, M.D. Eubanks, J.E. Losey and G.A. Langellotto. 1996. Habitat persistence underlies intraspecific variation in the dispersal strategies of planthoppers. Ecol. Monogr. 66: 389–408.CrossRefGoogle Scholar
  15. Dunson, W.A. and J. Travis. 1991. The role of abiotic factors in community organization. Am. Nat. 138: 1067–1091.CrossRefGoogle Scholar
  16. Fornara, D.A. and J.T. Du Toit. 2008. Browsing-induced effects on leaf litter quality and decomposition in a southern African savanna. Ecosystems 11: 238–249.CrossRefGoogle Scholar
  17. Gilliam, F.S. 2007. The ecological significance of the herbaceous layer in temperate forest ecosystems. Bioscience 57: 845–858.CrossRefGoogle Scholar
  18. Gomes da Silva, F.K., F. Lopes Sd, L.C. Serramo Lopez, J.I. Miranda de Melo and D.M. de Brito Melo Trovao. 2014. Patterns of species richness and conservation in the Caatinga along elevational gradients in a semiarid ecosystem. J. Arid. Environ. 110: 47–52.CrossRefGoogle Scholar
  19. Gray, A.N., T.A. Spies and M.J. Easter. 2002. Microclimatic and soil moisture responses to gap formation in coastal Douglas-fir forests. Can. J. Forest Res. 32: 332–343.CrossRefGoogle Scholar
  20. Grime, J.P. 1979. Plant Strategies and Vegetation Processes. John Wiley & Sons, Toronto.Google Scholar
  21. Grytnes, J.A. and O.R. Vetaas. 2002. Species richness and altitude: A comparison between null models and interpolated plant species richness along the Himalayan altitudinal gradient. Nepal. Am. Nat. 159: 294–304.CrossRefPubMedPubMedCentralGoogle Scholar
  22. Hill, J.K. and I.D. Hodkinson. 1995. Effects of temperature on phenological synchrony and altitudinal distribution of jumping plant lice (Hemiptera: Psylloidea) on dwarf willow (Salix lapponum) in Norway. Ecol. Entomol. 20: 237–244.CrossRefGoogle Scholar
  23. Hodkinson, I.D. 1999. Species response to global environmental change or why ecophysiological models are important: a reply to Davis et al. J. Anim. Ecol. 68: 1259-1262.CrossRefGoogle Scholar
  24. Hodkinson, I.D. 2005. Terrestrial insects along elevation gradients: species and community responses to altitude. Biol. Rev. 80: 489–513.CrossRefPubMedPubMedCentralGoogle Scholar
  25. Hutchinson, G.E. 1957. Concluding remarks. Population studies: animal ecology and demography. In: Cold Spring Harbor Symposia on Quantitative Biology. Cold Spring Harbor Lab Press. pp. 415– 427.Google Scholar
  26. Jeremy, M. and I.D. Hodkinson. 1999. Species at the edge of their range: the significance of the thermal environment for the distribution of congeneric Craspedolepta species (Sternorrhyncha: Psylloidea) living on Chamerion angustifolium (Onagraceae). Eur. J. Entomol. 96: 103–109.Google Scholar
  27. Kenward, M.G. and J.H. Roger. 1997. Small sample inference for fixed effects from restricted maximum likelihood. Biometrics 53: 983–997.CrossRefGoogle Scholar
  28. Kluge, J., M. Kessler and R.R. Dunn. 2006. What drives elevational patterns of diversity? A test of geometric constraints, climate and species pool effects for pteridophytes on an elevational gradient in Costa Rica. Global Ecol. Biogeogr. 15: 358–371.CrossRefGoogle Scholar
  29. Körner, C. and M. Diemer. 1994. Evidence that Plants from High Altitudes Retain their Greater Photosyntheti Effciency Under Elevated CO2. Funct. Ecol. 8: 58–68.CrossRefGoogle Scholar
  30. Körner, C. 2007. The use of ‘altitude’ in ecological research. Trends Ecol. Evol. 22: 569–574.CrossRefPubMedPubMedCentralGoogle Scholar
  31. Laliberté, E.B. Shipley and M.E. Laliberté. 2010. Package ‘FD’ Measuring functional diversity (FD) from multiple traits, and other tools for functional ecology.
  32. Legendre, P. and L. Legendre. 1998. Numerical Ecology. 2nd English edn. Elsevier, Amsterdam.Google Scholar
  33. Leuschner, C., G. Moser, C. Bertsch, M. Roederstein and D. Hertel. 2007. Large altitudinal increase in tree root/shoot ratio in tropical mountain forests of Ecuador. Basic Appl. Ecol. 8: 219–230.CrossRefGoogle Scholar
  34. Lowe, C.H. 1964. Arizona’s Natural Environment: Landscapes and Habitats. University of Arizona Press, Tucson, Arizona, USA.Google Scholar
  35. Ludwig, F., H. de Kroon, F. Berendse and H.H. Prins. 2004. The influence of savanna trees on nutrient, water and light availability and the understory vegetation. Plant Ecol. 170: 93–105.CrossRefGoogle Scholar
  36. Machac, A., M. Janda, R.R. Dunn and N.J. Sanders. 2011. Elevational gradients in phylogenetic structure of ant communities reveal the interplay of biotic and abiotic constraints on diversity. Ecography 34: 364–371.CrossRefGoogle Scholar
  37. Mölder, A., M. Bernhardt-Römermann and W. Schmidt. 2008. Herb-layer diversity in deciduous forests: raised by tree richness or beaten by beech? Forest. Ecol. Manag. 256: 272–281.CrossRefGoogle Scholar
  38. Mullah, C.J.A., K. Klanderud, O. Totland and B. Kigomo. 2014. Relationships between the density of two potential restoration tree species and plant species abundance and richness in a degraded Afromontane forest of Kenya. Afr. J. Ecol. 52:77–87.CrossRefGoogle Scholar
  39. Nelson, D.W. and L.A. Sommers. 1974. A rapid and accurate procedure for estimating organic carbon in soil. Proceedings of the Indiana Academy of Science pp. 456–462.Google Scholar
  40. Nogues-Bravo, D., M.B. Araujo, T. Lasanta and J.I. Lopez-Moreno. 2008. Climate change in Mediterranean mountains during the 21st century. Ambio 37: 280–285.CrossRefPubMedPubMedCentralGoogle Scholar
  41. Ohlemuller, R. and J.B. Wilson. 2000. Vascular plant species richness along latitudinal and altitudinal gradients: a contribution from New Zealand temperate rainforests. Ecol. Lett. 3: 262–266.CrossRefGoogle Scholar
  42. Oksanen, J.F.G. Blanchet, R. Kindt, M. J. Oksanen and M. Suggests. 2013. Package ‘vegan’ Community ecology package Version 2.3.
  43. Parmesan, C. 2006. Ecological and evolutionary responses to recent climate change. Annu. Rev. Ecol. Evol. S. 37: 637–669.CrossRefGoogle Scholar
  44. Price, J.N. and J.W. Morgan. 2008. Woody plant encroachment reduces species richness of herb-rich woodlands in southern Australia. Austral. Ecol. 33: 278–289.CrossRefGoogle Scholar
  45. Ramírez, J.M., P.J. Rey, J.M. Alcántara and A.M. Sánchez-Lafuente. 2006. Altitude and woody cover control recruitment of Helleborus foetidus in a Mediterranean mountain area. Ecography 29: 375–384.CrossRefGoogle Scholar
  46. Randall, M.G. 1982. The dynamics of an insect population throughout its altitudinal distribution: Coleophora alticolella (Lepidoptera) in northern England. J. Anim. Ecol. 51: 1: 993–1016.CrossRefGoogle Scholar
  47. Rochow, T.F. 1970. Ecological investigations of Thlaspi alpestre L. along an elevational gradient in Central Rocky Mountains. Ecology 51:649-&.CrossRefGoogle Scholar
  48. Roff, D. 1980. Optimizing development time in a seasonal environment: the ‘ups and downs’ of clinal variation. Oecologia 45: 202–208.CrossRefPubMedPubMedCentralGoogle Scholar
  49. Smith, K.J., W.S. Keeton, M.J. Twery and D.R. Tobi. 2008. Understory plant responses to uneven-aged forestry alternatives in northern hardwood-conifer forests. Can. J. Forest Res. 38: 1303–1318.CrossRefGoogle Scholar
  50. Sundqvist, M.K., N.J. Sanders and D.A. Wardle. 2013. Community and ecosystem responses to elevational gradients: processes, mechanisms, and insights for global change. Annu. Rev. Ecol. Evol. S. 44: 261–280.CrossRefGoogle Scholar
  51. Ter Steege, H. 1996. Winphot 5: a programme to analyze vegetation indices, light and light quality from hemispherical photographs. Tropenbos-Guyana Programme Tropenbos. Guyana.Google Scholar
  52. Vanhellemont, M., L. Baeten and K. Verheyen. 2014. Relating changes in understory diversity to environmental drivers in an ancient forest in northern Belgium. Plant Ecol. Evol. 147: 22–32.CrossRefGoogle Scholar
  53. Vourlitis, G.L., F.A. Lobo, S. Lawrence, K. Holt, A. Zappia, O.B. Pinto and J.D.S. Nogueira. 2014. Nutrient resorption in tropical savanna forests and woodlands of central Brazil. Plant Ecol. 215: 963–975.CrossRefGoogle Scholar
  54. Yun, F., M. Keming and Z. Yunxin. 2008. DCCA analysis of plant species distributions in different strata of oak (Quercus liaotungensis) forest along an altitudinal gradient in Dongling Mountain, China. J. Plant Ecol. (China Version). 32: 568–573.Google Scholar

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© Akadémiai Kiadó, Budapest 2015

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Authors and Affiliations

  • Zihan Jiang
    • 1
  • Keming Ma
    • 1
    • 3
    Email author
  • M. Anand
    • 2
  • Yuxin Zhang
    • 1
  1. 1.State Key Lab of Systems Ecology, Research Center for Eco-Environmental SciencesChinese Academy of SciencesBeijingPR China
  2. 2.School of Environmental SciencesUniv. of GuelphGuelphCanada
  3. 3.BeijingP. R. China

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