Late spring and late summer nonstructural carbohydrate (NSC) concentrations per unit of dry mass in leaves, aboveground woody tissues, and roots in large seedlings of shade tolerant, mid-shade tolerant, and shade intolerant species established in the understory of a cold rainforest and a Mediterranean forest in southern Chile. Bars represent mean values and error bars represent standard error; hatched and nonhatched areas symbolize starch and soluble sugars, respectively. Species included in each shade tolerance category are indicated in Table 1. Diff erent lower case letters and asterisks indicate statistically signifi cant diff erences at P < 0.05 between shade tolerance categories for a given sampling date and between dates for a given shade tolerance category, respectively, according to linear mixed-eff ect models. 

Contexts in source publication

Context 1

... Figs. 1, 2 ), which represents the most active period of growth in the studied species; i.e., when a storage-growth tradeoff should take place. Spring is also the pe- riod in which many evergreen species tend to experience their highest herbivory pressure ( Coley and Aide, 1991 ;Lowman, 1992 ;Carus, 2009 ), and hence, when traits (e.g., C storage) confer- ring advantages in the struggle against herbivores should be fully expressed. Th is result, then, is not consistent with the view that allocation of storage in woody tissues at the expense of growth promotes survival in low light conditions ( Kitajima, 1994 ;Kobe, 1997 ...

Context 2

... both forests, patterns of root and aboveground wood NSC and starch (but not of soluble sugars) in relation to shade tolerance were aff ected by sampling dates (i.e., signifi cant interaction Tolerance x Date, Table 2 , Figs. 1, 2 ) . However, late spring NSC and starch concentrations in woody tissues were not higher in shade tolerant species than in less tolerant species. In both forests, root NSC concentrations were similar between shade tolerant and intol- erant species in the late spring, and lower in shade tolerant than shade intolerant species in the late summer ( Fig. 1 ). In the cold rainforest, the latter was caused by a decrease in the NSC concen- tration of shade tolerant species in the late summer. In the Medi- terranean forest, in contrast, this pattern was driven by an increase in NSC and starch concentra- tions of shade intolerant species in the late summer ( Table 2 , Fig. 1 ). Furthermore, shade in- tolerant species in the Mediter- ranean forest increased their aboveground wood starch con- centrations in the late summer ( Table 2 , Fig. 1 ). As an excep- tion, aboveground wood NSC concentrations in the cold rain- forest were unaff ected by sam- pling date. Within a given shade toler- ance category, there was a high interspecifi c variation in NSC, starch, and soluble sugar con- centrations, and in the sampling date eff ects ( Fig. 3 ). Likewise, NSC fractions were very vari- able within tolerance categories, especially for roots and above- ground wood ( Fig. 4 ). Except for leaf NSC fractions, the interspe- cifi c variation in NSC was not apparently related to shade tol- erance ( Figs. 3, 4 ) ...

Context 3

... both forests, patterns of root and aboveground wood NSC and starch (but not of soluble sugars) in relation to shade tolerance were aff ected by sampling dates (i.e., signifi cant interaction Tolerance x Date, Table 2 , Figs. 1, 2 ) . However, late spring NSC and starch concentrations in woody tissues were not higher in shade tolerant species than in less tolerant species. In both forests, root NSC concentrations were similar between shade tolerant and intol- erant species in the late spring, and lower in shade tolerant than shade intolerant species in the late summer ( Fig. 1 ). In the cold rainforest, the latter was caused by a decrease in the NSC concen- tration of shade tolerant species in the late summer. In the Medi- terranean forest, in contrast, this pattern was driven by an increase in NSC and starch concentra- tions of shade intolerant species in the late summer ( Table 2 , Fig. 1 ). Furthermore, shade in- tolerant species in the Mediter- ranean forest increased their aboveground wood starch con- centrations in the late summer ( Table 2 , Fig. 1 ). As an excep- tion, aboveground wood NSC concentrations in the cold rain- forest were unaff ected by sam- pling date. Within a given shade toler- ance category, there was a high interspecifi c variation in NSC, starch, and soluble sugar con- centrations, and in the sampling date eff ects ( Fig. 3 ). Likewise, NSC fractions were very vari- able within tolerance categories, especially for roots and above- ground wood ( Fig. 4 ). Except for leaf NSC fractions, the interspe- cifi c variation in NSC was not apparently related to shade tol- erance ( Figs. 3, 4 ) ...

Context 4

... both forests, patterns of root and aboveground wood NSC and starch (but not of soluble sugars) in relation to shade tolerance were aff ected by sampling dates (i.e., signifi cant interaction Tolerance x Date, Table 2 , Figs. 1, 2 ) . However, late spring NSC and starch concentrations in woody tissues were not higher in shade tolerant species than in less tolerant species. In both forests, root NSC concentrations were similar between shade tolerant and intol- erant species in the late spring, and lower in shade tolerant than shade intolerant species in the late summer ( Fig. 1 ). In the cold rainforest, the latter was caused by a decrease in the NSC concen- tration of shade tolerant species in the late summer. In the Medi- terranean forest, in contrast, this pattern was driven by an increase in NSC and starch concentra- tions of shade intolerant species in the late summer ( Table 2 , Fig. 1 ). Furthermore, shade in- tolerant species in the Mediter- ranean forest increased their aboveground wood starch con- centrations in the late summer ( Table 2 , Fig. 1 ). As an excep- tion, aboveground wood NSC concentrations in the cold rain- forest were unaff ected by sam- pling date. Within a given shade toler- ance category, there was a high interspecifi c variation in NSC, starch, and soluble sugar con- centrations, and in the sampling date eff ects ( Fig. 3 ). Likewise, NSC fractions were very vari- able within tolerance categories, especially for roots and above- ground wood ( Fig. 4 ). Except for leaf NSC fractions, the interspe- cifi c variation in NSC was not apparently related to shade tol- erance ( Figs. 3, 4 ) ...

Context 5

... both forests, patterns of root and aboveground wood NSC and starch (but not of soluble sugars) in relation to shade tolerance were aff ected by sampling dates (i.e., signifi cant interaction Tolerance x Date, Table 2 , Figs. 1, 2 ) . However, late spring NSC and starch concentrations in woody tissues were not higher in shade tolerant species than in less tolerant species. In both forests, root NSC concentrations were similar between shade tolerant and intol- erant species in the late spring, and lower in shade tolerant than shade intolerant species in the late summer ( Fig. 1 ). In the cold rainforest, the latter was caused by a decrease in the NSC concen- tration of shade tolerant species in the late summer. In the Medi- terranean forest, in contrast, this pattern was driven by an increase in NSC and starch concentra- tions of shade intolerant species in the late summer ( Table 2 , Fig. 1 ). Furthermore, shade in- tolerant species in the Mediter- ranean forest increased their aboveground wood starch con- centrations in the late summer ( Table 2 , Fig. 1 ). As an excep- tion, aboveground wood NSC concentrations in the cold rain- forest were unaff ected by sam- pling date. Within a given shade toler- ance category, there was a high interspecifi c variation in NSC, starch, and soluble sugar con- centrations, and in the sampling date eff ects ( Fig. 3 ). Likewise, NSC fractions were very vari- able within tolerance categories, especially for roots and above- ground wood ( Fig. 4 ). Except for leaf NSC fractions, the interspe- cifi c variation in NSC was not apparently related to shade tol- erance ( Figs. 3, 4 ) ...

Context 6

... e lack of relationship between root and aboveground NSC in the late spring and shade tolerance could also occur if C storage pat- terns were driven by drought tolerance or fi re resistance ( Poorter and Kitajima, 2007 ;Poorter et al., 2010 ). Indeed, C storage prolongs plant survival under drought by mediating osmoregulation and mainte- nance of vascular integrity ( Hartmann et al., 2013 ;Mitchell et al., 2013 ;' O'Brien et al., 2014 ), and is important for resprouting aft er fi re ( Bond and Midgley, 2001 ). High levels of C storage could be advanta- geous to recover from drought and fi re, which are main causes of seedling mortality in Mediterranean forests. Furthermore, drought and shade tolerance were found to be unrelated in Mediterranean for- ests ( S á nchez- Gómez et al., 2006 ), even with similar climate and physiognomy to the one I studied ( Parada and Lusk, 2011 ). Likewise, shade tolerance and fi re resistance are not apparently related in the Mediterranean species studied here, because resprouting has been reported in both highly shade tolerant and intolerant species ( Veblen et al., 1979 ;Gonz á lez et al., 2002 ;Donoso, 2006 ). Although the cold rainforest is characterized by high rainfall, the root and aboveground NSC patterns in relation to shade tolerance at this site could also have Late spring and late summer NSC concentrations per unit of dry mass in leaves, aboveground woody tissues, and roots in seedlings of species diff ering in light requirements, established in the understory of a cold rainforest and a Mediterranean forest in Chile. Bars represent mean values and error bars represent standard error; hatched and nonhatched areas symbolize starch and soluble sugar concentrations, respectively. The x -axis indicates abbreviated species names according to Table 1 . been driven by drought. In very humid tropical forests, drought is a major determinant of seed- ling mortality and species distri- bution ( Engelbrecht et al., 2005 ), and NSC concentrations were found to positively correlate with drought tolerance in seed- lings ( O'Brien et al., 2014 ). It has been suggested that a drought- shade tolerance tradeoff under- lies southern Chilean rainforests ( Jiménez-Castillo et al., 2011 ); this could explain the higher aboveground wood NSC con- centration in shade intolerant species compared to tolerant species in the cold rainforest during late spring. Th e proposed idea that drought may cause in- terspecifi c variation in C storage in both forests is supported by the fact that diff erences in root and aboveground wood NSC concentrations among shade tol- erance categories were caused by diff erences in soluble sugars (i.e., osmotically active) rather than starch ( Table 2 , Fig. 1 ...

Context 7

... e sampling date had substantial eff ects on the NSC and starch concentration patterns in woody tissues in relation to shade tolerance, revealing seasonal inconsistency in the relationship between C storage and shade tolerance. In addition, such inconsistencies were diff erent between forests. For example, shade tolerant species of the cold rainfor- est decreased their root NSC and starch concentrations in summer, but this was not observed in the Mediterranean forest ( Fig. 1 ). Likewise, FIGURE 4 Late spring and late summer fractions of NSC pool in leaves, aboveground woody tissues, and roots in seedlings of species diff ering in light requirements, established in the understory of a cold rainforest and a Mediterranean forest in Chile. Bars represent mean values and error bars represent standard error. The x -axis indicates abbreviated species names according to Table 1 . shade intolerant species of the Mediterranean forest increased their root NSC concentrations in late summer; this was not observed in the cold rainforest ( Table 2 , Fig. 1 ). Decreasing NSC concen- trations during the growing season are common in evergreen species ( Hoch et al., 2003 ;Rosas et al., 2013 ), and the lack of such decrease in shade intolerant and mid-tolerant species of the cold rainforest could be related to an- ticipated freezing temperatures af- fecting the C sinks (e.g., respiration, bud formation) more than photo- synthesis. Increased late summer NSC concentrations in shade intol- erant species of the Mediterranean forest could result from an adaptive strategy to provide carbohydrates under drought ( Dietze et al., 2014 ), or from higher photosynthetic rates when the winter deciduous overstory opened in the late sum- mer. Regardless of the mechanisms involved, this result implies that potential storage-growth tradeoff s (underlying mechanisms of shade tolerance) cannot be inferred by assessing C storage at the end of the growing ...

Context 8

... e sampling date had substantial eff ects on the NSC and starch concentration patterns in woody tissues in relation to shade tolerance, revealing seasonal inconsistency in the relationship between C storage and shade tolerance. In addition, such inconsistencies were diff erent between forests. For example, shade tolerant species of the cold rainfor- est decreased their root NSC and starch concentrations in summer, but this was not observed in the Mediterranean forest ( Fig. 1 ). Likewise, FIGURE 4 Late spring and late summer fractions of NSC pool in leaves, aboveground woody tissues, and roots in seedlings of species diff ering in light requirements, established in the understory of a cold rainforest and a Mediterranean forest in Chile. Bars represent mean values and error bars represent standard error. The x -axis indicates abbreviated species names according to Table 1 . shade intolerant species of the Mediterranean forest increased their root NSC concentrations in late summer; this was not observed in the cold rainforest ( Table 2 , Fig. 1 ). Decreasing NSC concen- trations during the growing season are common in evergreen species ( Hoch et al., 2003 ;Rosas et al., 2013 ), and the lack of such decrease in shade intolerant and mid-tolerant species of the cold rainforest could be related to an- ticipated freezing temperatures af- fecting the C sinks (e.g., respiration, bud formation) more than photo- synthesis. Increased late summer NSC concentrations in shade intol- erant species of the Mediterranean forest could result from an adaptive strategy to provide carbohydrates under drought ( Dietze et al., 2014 ), or from higher photosynthetic rates when the winter deciduous overstory opened in the late sum- mer. Regardless of the mechanisms involved, this result implies that potential storage-growth tradeoff s (underlying mechanisms of shade tolerance) cannot be inferred by assessing C storage at the end of the growing ...

Context 9

... Figs. 1, 2 ), which represents the most active period of growth in the studied species; i.e., when a storage-growth tradeoff should take place. Spring is also the pe- riod in which many evergreen species tend to experience their highest herbivory pressure ( Coley and Aide, 1991 ;Lowman, 1992 ;Carus, 2009 ), and hence, when traits (e.g., C storage) confer- ring advantages in the struggle against herbivores should be fully expressed. Th is result, then, is not consistent with the view that allocation of storage in woody tissues at the expense of growth promotes survival in low light conditions ( Kitajima, 1994 ;Kobe, 1997 ...

Context 10

... both forests, patterns of root and aboveground wood NSC and starch (but not of soluble sugars) in relation to shade tolerance were aff ected by sampling dates (i.e., signifi cant interaction Tolerance x Date, Table 2 , Figs. 1, 2 ) . However, late spring NSC and starch concentrations in woody tissues were not higher in shade tolerant species than in less tolerant species. In both forests, root NSC concentrations were similar between shade tolerant and intol- erant species in the late spring, and lower in shade tolerant than shade intolerant species in the late summer ( Fig. 1 ). In the cold rainforest, the latter was caused by a decrease in the NSC concen- tration of shade tolerant species in the late summer. In the Medi- terranean forest, in contrast, this pattern was driven by an increase in NSC and starch concentra- tions of shade intolerant species in the late summer ( Table 2 , Fig. 1 ). Furthermore, shade in- tolerant species in the Mediter- ranean forest increased their aboveground wood starch con- centrations in the late summer ( Table 2 , Fig. 1 ). As an excep- tion, aboveground wood NSC concentrations in the cold rain- forest were unaff ected by sam- pling date. Within a given shade toler- ance category, there was a high interspecifi c variation in NSC, starch, and soluble sugar con- centrations, and in the sampling date eff ects ( Fig. 3 ). Likewise, NSC fractions were very vari- able within tolerance categories, especially for roots and above- ground wood ( Fig. 4 ). Except for leaf NSC fractions, the interspe- cifi c variation in NSC was not apparently related to shade tol- erance ( Figs. 3, 4 ) ...

Context 11

... both forests, patterns of root and aboveground wood NSC and starch (but not of soluble sugars) in relation to shade tolerance were aff ected by sampling dates (i.e., signifi cant interaction Tolerance x Date, Table 2 , Figs. 1, 2 ) . However, late spring NSC and starch concentrations in woody tissues were not higher in shade tolerant species than in less tolerant species. In both forests, root NSC concentrations were similar between shade tolerant and intol- erant species in the late spring, and lower in shade tolerant than shade intolerant species in the late summer ( Fig. 1 ). In the cold rainforest, the latter was caused by a decrease in the NSC concen- tration of shade tolerant species in the late summer. In the Medi- terranean forest, in contrast, this pattern was driven by an increase in NSC and starch concentra- tions of shade intolerant species in the late summer ( Table 2 , Fig. 1 ). Furthermore, shade in- tolerant species in the Mediter- ranean forest increased their aboveground wood starch con- centrations in the late summer ( Table 2 , Fig. 1 ). As an excep- tion, aboveground wood NSC concentrations in the cold rain- forest were unaff ected by sam- pling date. Within a given shade toler- ance category, there was a high interspecifi c variation in NSC, starch, and soluble sugar con- centrations, and in the sampling date eff ects ( Fig. 3 ). Likewise, NSC fractions were very vari- able within tolerance categories, especially for roots and above- ground wood ( Fig. 4 ). Except for leaf NSC fractions, the interspe- cifi c variation in NSC was not apparently related to shade tol- erance ( Figs. 3, 4 ) ...

Context 12

... both forests, patterns of root and aboveground wood NSC and starch (but not of soluble sugars) in relation to shade tolerance were aff ected by sampling dates (i.e., signifi cant interaction Tolerance x Date, Table 2 , Figs. 1, 2 ) . However, late spring NSC and starch concentrations in woody tissues were not higher in shade tolerant species than in less tolerant species. In both forests, root NSC concentrations were similar between shade tolerant and intol- erant species in the late spring, and lower in shade tolerant than shade intolerant species in the late summer ( Fig. 1 ). In the cold rainforest, the latter was caused by a decrease in the NSC concen- tration of shade tolerant species in the late summer. In the Medi- terranean forest, in contrast, this pattern was driven by an increase in NSC and starch concentra- tions of shade intolerant species in the late summer ( Table 2 , Fig. 1 ). Furthermore, shade in- tolerant species in the Mediter- ranean forest increased their aboveground wood starch con- centrations in the late summer ( Table 2 , Fig. 1 ). As an excep- tion, aboveground wood NSC concentrations in the cold rain- forest were unaff ected by sam- pling date. Within a given shade toler- ance category, there was a high interspecifi c variation in NSC, starch, and soluble sugar con- centrations, and in the sampling date eff ects ( Fig. 3 ). Likewise, NSC fractions were very vari- able within tolerance categories, especially for roots and above- ground wood ( Fig. 4 ). Except for leaf NSC fractions, the interspe- cifi c variation in NSC was not apparently related to shade tol- erance ( Figs. 3, 4 ) ...

Context 13

... both forests, patterns of root and aboveground wood NSC and starch (but not of soluble sugars) in relation to shade tolerance were aff ected by sampling dates (i.e., signifi cant interaction Tolerance x Date, Table 2 , Figs. 1, 2 ) . However, late spring NSC and starch concentrations in woody tissues were not higher in shade tolerant species than in less tolerant species. In both forests, root NSC concentrations were similar between shade tolerant and intol- erant species in the late spring, and lower in shade tolerant than shade intolerant species in the late summer ( Fig. 1 ). In the cold rainforest, the latter was caused by a decrease in the NSC concen- tration of shade tolerant species in the late summer. In the Medi- terranean forest, in contrast, this pattern was driven by an increase in NSC and starch concentra- tions of shade intolerant species in the late summer ( Table 2 , Fig. 1 ). Furthermore, shade in- tolerant species in the Mediter- ranean forest increased their aboveground wood starch con- centrations in the late summer ( Table 2 , Fig. 1 ). As an excep- tion, aboveground wood NSC concentrations in the cold rain- forest were unaff ected by sam- pling date. Within a given shade toler- ance category, there was a high interspecifi c variation in NSC, starch, and soluble sugar con- centrations, and in the sampling date eff ects ( Fig. 3 ). Likewise, NSC fractions were very vari- able within tolerance categories, especially for roots and above- ground wood ( Fig. 4 ). Except for leaf NSC fractions, the interspe- cifi c variation in NSC was not apparently related to shade tol- erance ( Figs. 3, 4 ) ...

Context 14

... e lack of relationship between root and aboveground NSC in the late spring and shade tolerance could also occur if C storage pat- terns were driven by drought tolerance or fi re resistance ( Poorter and Kitajima, 2007 ;Poorter et al., 2010 ). Indeed, C storage prolongs plant survival under drought by mediating osmoregulation and mainte- nance of vascular integrity ( Hartmann et al., 2013 ;Mitchell et al., 2013 ;' O'Brien et al., 2014 ), and is important for resprouting aft er fi re ( Bond and Midgley, 2001 ). High levels of C storage could be advanta- geous to recover from drought and fi re, which are main causes of seedling mortality in Mediterranean forests. Furthermore, drought and shade tolerance were found to be unrelated in Mediterranean for- ests ( S á nchez- Gómez et al., 2006 ), even with similar climate and physiognomy to the one I studied ( Parada and Lusk, 2011 ). Likewise, shade tolerance and fi re resistance are not apparently related in the Mediterranean species studied here, because resprouting has been reported in both highly shade tolerant and intolerant species ( Veblen et al., 1979 ;Gonz á lez et al., 2002 ;Donoso, 2006 ). Although the cold rainforest is characterized by high rainfall, the root and aboveground NSC patterns in relation to shade tolerance at this site could also have Late spring and late summer NSC concentrations per unit of dry mass in leaves, aboveground woody tissues, and roots in seedlings of species diff ering in light requirements, established in the understory of a cold rainforest and a Mediterranean forest in Chile. Bars represent mean values and error bars represent standard error; hatched and nonhatched areas symbolize starch and soluble sugar concentrations, respectively. The x -axis indicates abbreviated species names according to Table 1 . been driven by drought. In very humid tropical forests, drought is a major determinant of seed- ling mortality and species distri- bution ( Engelbrecht et al., 2005 ), and NSC concentrations were found to positively correlate with drought tolerance in seed- lings ( O'Brien et al., 2014 ). It has been suggested that a drought- shade tolerance tradeoff under- lies southern Chilean rainforests ( Jiménez-Castillo et al., 2011 ); this could explain the higher aboveground wood NSC con- centration in shade intolerant species compared to tolerant species in the cold rainforest during late spring. Th e proposed idea that drought may cause in- terspecifi c variation in C storage in both forests is supported by the fact that diff erences in root and aboveground wood NSC concentrations among shade tol- erance categories were caused by diff erences in soluble sugars (i.e., osmotically active) rather than starch ( Table 2 , Fig. 1 ...

Context 15

... e sampling date had substantial eff ects on the NSC and starch concentration patterns in woody tissues in relation to shade tolerance, revealing seasonal inconsistency in the relationship between C storage and shade tolerance. In addition, such inconsistencies were diff erent between forests. For example, shade tolerant species of the cold rainfor- est decreased their root NSC and starch concentrations in summer, but this was not observed in the Mediterranean forest ( Fig. 1 ). Likewise, FIGURE 4 Late spring and late summer fractions of NSC pool in leaves, aboveground woody tissues, and roots in seedlings of species diff ering in light requirements, established in the understory of a cold rainforest and a Mediterranean forest in Chile. Bars represent mean values and error bars represent standard error. The x -axis indicates abbreviated species names according to Table 1 . shade intolerant species of the Mediterranean forest increased their root NSC concentrations in late summer; this was not observed in the cold rainforest ( Table 2 , Fig. 1 ). Decreasing NSC concen- trations during the growing season are common in evergreen species ( Hoch et al., 2003 ;Rosas et al., 2013 ), and the lack of such decrease in shade intolerant and mid-tolerant species of the cold rainforest could be related to an- ticipated freezing temperatures af- fecting the C sinks (e.g., respiration, bud formation) more than photo- synthesis. Increased late summer NSC concentrations in shade intol- erant species of the Mediterranean forest could result from an adaptive strategy to provide carbohydrates under drought ( Dietze et al., 2014 ), or from higher photosynthetic rates when the winter deciduous overstory opened in the late sum- mer. Regardless of the mechanisms involved, this result implies that potential storage-growth tradeoff s (underlying mechanisms of shade tolerance) cannot be inferred by assessing C storage at the end of the growing ...

Context 16

... e sampling date had substantial eff ects on the NSC and starch concentration patterns in woody tissues in relation to shade tolerance, revealing seasonal inconsistency in the relationship between C storage and shade tolerance. In addition, such inconsistencies were diff erent between forests. For example, shade tolerant species of the cold rainfor- est decreased their root NSC and starch concentrations in summer, but this was not observed in the Mediterranean forest ( Fig. 1 ). Likewise, FIGURE 4 Late spring and late summer fractions of NSC pool in leaves, aboveground woody tissues, and roots in seedlings of species diff ering in light requirements, established in the understory of a cold rainforest and a Mediterranean forest in Chile. Bars represent mean values and error bars represent standard error. The x -axis indicates abbreviated species names according to Table 1 . shade intolerant species of the Mediterranean forest increased their root NSC concentrations in late summer; this was not observed in the cold rainforest ( Table 2 , Fig. 1 ). Decreasing NSC concen- trations during the growing season are common in evergreen species ( Hoch et al., 2003 ;Rosas et al., 2013 ), and the lack of such decrease in shade intolerant and mid-tolerant species of the cold rainforest could be related to an- ticipated freezing temperatures af- fecting the C sinks (e.g., respiration, bud formation) more than photo- synthesis. Increased late summer NSC concentrations in shade intol- erant species of the Mediterranean forest could result from an adaptive strategy to provide carbohydrates under drought ( Dietze et al., 2014 ), or from higher photosynthetic rates when the winter deciduous overstory opened in the late sum- mer. Regardless of the mechanisms involved, this result implies that potential storage-growth tradeoff s (underlying mechanisms of shade tolerance) cannot be inferred by assessing C storage at the end of the growing ...