Table 3.

Results of nested ANOVA testing for differences among leaf traits on a mass-basis between sun and shade leaves and site¹.

Response variable	Source of variation
	Site		Canopy position (site)		Offset
	F	P	F	P	µ ± STERR
Light capture and growth
LMA	22.97	<0.01	7.70	<0.01	−19.18 ± 1.04*
N	31.63	<0.01	0.30	NS
Chlorophyll ab	19.26	<0.01	8.33	<0.01	1.43 ± 0.10*
Carotenoids	19.01	<0.01	4.35	<0.01	0.21 ± 0.02*
Soluble C	30.90	<0.01	0.35	NS
δ13C	35.22	<0.01	16.56	<0.01	−1.42 ± 0.06*
Structure and defense
Total C	23.98	<0.01	0.75	NS
Lignin	11.34	<0.01	0.26	NS
Cellulose	28.65	<0.01	1.46	NS
Phenols	24.58	<0.01	0.30	NS
Tannins	37.85	<0.01	0.20	NS
Macronutrients
P	36.95	<0.01	0.35	NS
Ca	132.97	<0.01	0.46	NS
K	11.93	<0.01	1.70	NS
Mg	25.31	<0.01	0.61	NS
Micronutrients
B	23.99	<0.01	0.24	NS
Fe	23.39	<0.01	1.63	NS
Mn	68.02	<0.01	0.08	NS
Zn	5.01	<0.01	0.52	NS

¹Sample collection at sites varying in elevation, light environment, and geology meant that canopy position was effectively nested within site.

LMA, leaf mass per unit area; LMA, N, chlorophyll ab, carotenoids, P, K, and Mg were log-transformed before analysis. Ca was transformed by square root.

STERR is standard error.

* indicates significant offset at p < 0.01. NS is nonsignificant.

Mean offset among canopy position (shade to sun) across all sites is also shown (Matched pairs t-test, p <0.001). LMA is leaf mass per area. NSC are nonstructural carbohydrates.