Ecological Research

, Volume 29, Issue 4, pp 607–617 | Cite as

Leaf-trait responses to environmental gradients in moorland communities: contribution of intraspecific variation, species replacement and functional group replacement

  • Chiho Kamiyama
  • Masatoshi Katabuchi
  • Takehiro Sasaki
  • Masaya Shimazaki
  • Tohru Nakashizuka
  • Kouki Hikosaka
Original Article


The values of many important traits of plants in a community change along environmental gradients. Such changes may involve intraspecific variation and replacement by species that have different trait values. We hypothesized that they also involve the variation within and among functional groups (FGs) to the environmental dependence of trait values at the community level. We studied environmental dependence of trait values in 27 moorlands at various scales and analyzed to what extent intraspecific variation, species replacement within FGs and FG replacement contribute to the gradient of community trait values. The community structure in moorlands was influenced mainly by two environmental factors: temperature and water condition. Plants inhabiting sites with low temperature and low-pH generally tended to have lower maximum leaf height, greater leaf mass per area, and smaller leaf size. At the community level, site-mean of maximum leaf height and leaf size generally increased with increasing temperature and water pH. Our analysis demonstrated that the relative contributions of intraspecific variation, species replacement within FGs and FG replacement differed depending on combinations of the traits and environments. The contribution of FG replacement varied considerably among cases (0.6–34.5 %). Species replacement within FGs, which has received little attention in previous studies, was most responsible for the community-level changes (31.6–65.3 %) and intraspecific variation also made a large contribution (22.9–57.9 %). Understanding such various mechanisms involving intraspecific variation and species replacement should help us better predict how the structure and functioning of moorland plant communities will respond to climate change.


Functional diversity Functional traits Growth form Leaf habit Altitude Leaf height Leaf mass area Leaf size 



We thank the lab members for their kindness in helping with the field work, especially Yukari Abe, Sawako Imahiro, Akira Inoue, and Yuta Nihonmatsu. We also thank Koji Yonekura for his advice about species identification and Hiroko Kurokawa for her helpful comments. The study was conducted with permission by the Ministry of the Environment, Aomori Prefecture Government, Aomori City and a landowner, Tashiro farm. This work was carried out under a JSPS research fellowship awarded to CK, and supported by KAKENHI to KH, the Global Environment Research Fund (F-052 and F-092) by the Ministry of the Environment, Japan to KH, a research grant from The Mitsui & Co., Ltd. Environment Fund, and the Global COE Program “Ecosystem Adaptability Science for the Future” (j03) of the MEXT.

Supplementary material

11284_2014_1148_MOESM1_ESM.doc (554 kb)
Supplementary material 1 (DOC 554 kb)


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Copyright information

© The Ecological Society of Japan 2014

Authors and Affiliations

  • Chiho Kamiyama
    • 1
    • 2
  • Masatoshi Katabuchi
    • 3
  • Takehiro Sasaki
    • 4
  • Masaya Shimazaki
    • 2
  • Tohru Nakashizuka
    • 2
  • Kouki Hikosaka
    • 2
  1. 1.Institute for the Advanced Study of SustainabilityUnited Nations UniversityShibuyaJapan
  2. 2.Graduate School of Life SciencesTohoku UniversitySendaiJapan
  3. 3.Key Laboratory of Tropical Forest Ecology, Xishuangbanna Tropical Botanical GardenChinese Academy of SciencesMenglaChina
  4. 4.Graduate School of Frontier SciencesThe University of TokyoKashiwaJapan

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