Leaf-trait responses to environmental gradients in moorland communities: contribution of intraspecific variation, species replacement and functional group replacement
- 455 Downloads
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.
KeywordsFunctional 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.
- Brady N, Weil R (1998) The nature and properties of soils, 12th edn. Macmillan, New YorkGoogle Scholar
- Diaz S, Hodgson JG, Thompson K, Cabido M, Cornelissen JHC, Jalili A, Montserrat-Martí G, Grime JP, Zarrinkamar F, Asri Y, Band SR, Basconcelo S, Castro-Díez P, Funes G, Hamzehee B, Khoshnevi M, Pérez-Harguindeguy N, Pérez-Rontomé MC, Shirvany FA, Vendramini F, Yazdani S, Abbas-Azimi R, Bogaard A, Boustani S, Charles M, Dehghan M, de Torres-Espuny L, Falczuk V, Guerrero-Campo J, Hynd A, Jones G, Kowsary E, Kazemi-Saeed F, Maestro-Martínez M, Romo-Díez A, Shaw S, Siavash B, Villar-Salvador P, Zak MR (2004) The plant traits that drive ecosystems: evidence from three continents. J Veg Sci 15:295–304CrossRefGoogle Scholar
- Givnish T (2002) Adaptive significance of evergreen vs. deciduous leaves: solving the triple paradox. Silva Fenn 36:703–743Google Scholar
- Koike K, Toshikazu T, Chinzei K, Miyagi T (2005) Regional geomorphology of the Japanese Islands, vol 3. Geomorphology of Tohoku Region. University of Tokyo Press, Tokyo (in Japanese)Google Scholar
- Körner C, Neumayer M, Pelaez Mennendez-Riedl S, Smeets-Scheel A (1989) Functional morphology of mountain plants. Flora 182:353–383Google Scholar
- Mochida Y, Sato M, Kikuchi T (1997) The vegetation of the south Hakkoda Mountains, northern Honshu. Jpn Ecol Rev 23:295–300Google Scholar
- Muraoka H, Takakura S (1988) Explanatory text of the geological map of the Hakkôda Geothermal Area (in Japanese). Geological Survey of Japan, TsukubaGoogle Scholar
- Nagano S, Nakano T, Hikosaka K, Maruta E (2009) Needle traits of an evergreen, coniferous shrub growing at wind-exposed and protected sites in a mountain region: does Pinus pumila produce needles with greater mass per area under wind-stress conditions? Plant Biol 11:94–100PubMedCrossRefGoogle Scholar
- Schimper A (1903) Plant-geography upon a physiological basis. Clarendon Press, OxfordGoogle Scholar
- Spasojevic MJ, Suding KN (2012) Inferring community assembly mechanisms from functional diversity patterns: the importance of multiple assembly processes. J Ecol 100:652–661Google Scholar
- Wright IJ, Reich PB, Westoby M, Ackerly DD, Baruch Z, Bongers F, Cavender-Bares J, Chapin T, Cornelissen JHC, Diemer M, Flexas J, Garnier E, Groom PK, Gulias J, Hikosaka K, Lamont BB, Lee T, Lee W, Lusk C, Midgley JJ, Navas M-L, Niinemets U, Oleksyn J, Osada N, Poorter H, Poot P, Prior L, Pyankov VI, Roumet C, Thomas SC, Tjoelker MG, Veneklaas EJ, Villar R (2004) The worldwide leaf economics spectrum. Nature 428:821–827PubMedCrossRefGoogle Scholar