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Box 9E. 1 Continued FIGURE 2. The C-S-R triangle model (Grime
1979). The strategies at the three corners are C, competiti-
winning species; S, stress-tolerating s- cies; R,ruderalspecies.
Particular species can engage in any mixture of these three primary
strategies, and the m- ture is described by their position within
the triangle. comment briefly on some other dimensions that Grime's
(1977) triangle (Fig. 2) (see also Sects. 6. 1 are not yet so well
understood. and 6. 3 of Chapter 7 on growth and allocation) is a
two-dimensional scheme. A C-S axis (Com- tition-winning species to
Stress-tolerating spe- Leaf Economics Spectrum cies) reflects
adaptation to favorable vs. unfavorable sites for plant growth, and
an R- Five traits that are coordinated across species are axis
(Ruderal species) reflects adaptation to leaf mass per area (LMA),
leaf life-span, leaf N disturbance. concentration, and potential
photosynthesis and dark respiration on a mass basis. In the
five-trait Trait-Dimensions space,79%ofallvariation
worldwideliesalonga single main axis (Fig. 33 of Chapter 2A on
photo- A recent trend in plant strategy thinking has synthesis;
Wright et al. 2004). Species with low been trait-dimensions, that
is, spectra of varia- LMA tend to have short leaf life-spans, high
leaf tion with respect to measurable traits. Compared nutrient
concentrations, and high potential rates of mass-based
photosynthesis. These species with category schemes, such as
Raunkiaer's, trait occur at the ''quick-return'' end of the leaf e-
dimensions have the merit of capturing cont- nomics spectrum.
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