# Climatic Factors Determine the Distribution Patterns of Leaf Nutrient Traits at Large Scales

^{1}

^{2}

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## Abstract

**:**

## 1. Introduction

## 2. Results

#### 2.1. Variability in LN, LP, and N/P of Different Life Forms and Distribution Patterns in China

#### 2.2. Correlations between Climatic Factors and LN, LP, and N/P at Different Life Forms

^{2}= 0.20, p < 0.001; R

^{2}= 0.16, p < 0.001) and better predictive power for LP in the tree layer (R

^{2}= 0.46, p < 0.001; R

^{2}= 0.50, p < 0.001; R

^{2}= 0.37, p < 0.001) were MWMT, MAP, and ASD.

^{2}= 0.59, p < 0.001).

#### 2.3. Effect of Soil Factors on the Relationship between LN, LP, and Leaf N/P at Different Life Forms

^{2}= 0.30, p < 0.001) (Figure 8a). Soil pH (Figure 7c) was the best predictor of LP in the herb and tree levels (R

^{2}= 0.25, p < 0.001; R

^{2}= 0.29, p < 0.001) (Figure 7c), and the best predictor of N/P in the tree level (Figure 8c) (R

^{2}= 0.29, p < 0.001) (Figure 8c). In contrast, the best prediction of N/P for the shrub level was for soil P (R

^{2}= 0.22, p < 0.001; Figure 8b).

#### 2.4. Climatic and Soil Factors Dominate Changes in the Functional Traits of Different Communities

## 3. Discussion

## 4. Materials and Methods

#### 4.1. Study Area and Sample Data

#### 4.2. Functional Data

_{i}) represents the forest mean trait values.

_{i}represents the community−weighted functional trait identity value and D

_{i}represents the abundance of dominant tree species. Trait

_{i}represents the selected functional trait [50].

#### 4.3. Environmental Data

#### 4.4. Data Analysis

^{2}represents how well the model fits the variables studied.

_{i}is the explanatory variable that strictly follows the parametric form, β

_{i}is the corresponding parameter, and f

_{j}($\mathrm{X}$

_{j}) is the corresponding explanatory variable that follows the nonparametric form of the smoothing function. In our study, the spline smoothing function S(•) was selected for fitting, thin−plate spline smoothing was selected for function fitting between different nodes, and each smoothing function S(•) was estimated using penalized least squares [50].

## 5. Conclusions

## Supplementary Materials

## Author Contributions

## Funding

## Institutional Review Board Statement

## Informed Consent Statement

## Data Availability Statement

## Conflicts of Interest

## References

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**Figure 1.**The distribution patterns of LN, LP, and N/P at different life forms in China with a spatial resolution of 1 × 1 km were studied by kernel density estimation. (

**a**) LN of the tree levels; (

**b**) LP of the tree levels; (

**c**) N/P of the tree levels; (

**d**) LN of the shrub levels; (

**e**) LP of the shrub levels; (

**f**) leaf N/P of the shrub levels; (

**g**) LN of the herb levels; (

**h**) herb levels LP; (

**i**) herb levels N/P.

**Figure 2.**A comparison of the differences in the LN (

**a**), LP (

**b**), and N/P (

**c**) at different life forms. (

**a**) Variability of LN among the trees, shrubs, and herbs; (

**b**) variability of LP among the trees, shrubs, and herbs; (

**c**) variability of N/P among the trees, shrubs, and herbs. Tree represents the tree levels, Shrub represents the shrub levels, and Herb represents the herb levels. Levels are grouped where ns represents non−significant (p > 0.05) at the 0.05 level, * represents p < 0.05, ** represents p < 0.01, **** represents p < 0.0001.

**Figure 3.**The general linear correlation analysis of climate factors with LN at different life forms. (

**a**) General linear relationship between the MAT and LN of plants in the trees, shrubs, and herbs. (

**b**) General linear relationship between the MCMT and LN of plants in the trees, shrubs, and herbs. (

**c**) General linear relationship between the MWMT and LN of plants in the trees, shrubs, and herbs (

**d**) General linear relationship between the MAP and LN of plants in the trees, shrubs, and herbs (

**e**) General linear relationship between the ASD and LN of plants in the trees, shrubs, and herbs (

**f**) General linear relationship between the MAE and LN of plants in the trees, shrubs, and herbs. MAT represents the mean annual temperature, MCMT represents the mean coldest monthly temperature, MWMT represents the mean warmest monthly temperature, MAP represents the mean annual precipitation, ASD represents the annual sunlight duration, and MAE represents the mean annual evaporation. The red line represents the tree level, the green line is the shrub level, and the blue line is the herb level. R

^{2}represents how well the model fits the variables studied and the p-value represents the significance level.

**Figure 4.**The general linear correlation analysis of climate factors with LP at different life forms. (

**a**) General linear relationship between the MAT and LP of plants in the trees, shrubs, and herbs. (

**b**) General linear relationship between the MCMT and LP of plants in the trees, shrubs, and herbs. (

**c**) General linear relationship between the MWMT and LP of plants in the trees, shrubs, and herbs. (

**d**) General linear relationship between the MAP and LP of plants in the trees, shrubs, and herbs. (

**e**) General linear relationship between the ASD and LP of plants in the trees, shrubs, and herbs. (

**f**) General linear relationship between the MAE and LP of plants in the trees, shrubs, and herbs. MAT represents the mean annual temperature, MCMT represents the mean coldest monthly temperature, MWMT represents the mean warmest monthly temperature, MAP represents the mean annual precipitation, ASD represents the annual sunlight duration, and MAE represents the mean annual evaporation. The red line represents the tree level, the green line is the shrub level, and the blue line is the herb level. R

^{2}represents how well the model fits the variables studied and the p-value represents the significance level.

**Figure 5.**The general linear correlation analysis of climate factors with N/P at different life forms. (

**a**) General linear relationship between the MAT and N/P of plants in the trees, shrubs, and herbs. (

**b**) General linear relationship between the MCMT and N/P of plants in the trees, shrubs, and herbs. (

**c**) General linear relationship between the MWMT and N/P of plants in the trees, shrubs, and herbs. (

**d**) General linear relationship between the MAP and N/P of plants in the trees, shrubs, and herbs. (

**e**) General linear relationship between the ASD and N/P of plants in the trees, shrubs, and herbs. (

**f**): General linear relationship between the MAE and N/P of plants in the trees, shrubs, and herbs. MAT represents the mean annual temperature, MCMT represents the mean coldest monthly temperature, MWMT represents the mean warmest monthly temperature, MAP represents the mean annual precipitation, ASD represents the annual sunlight duration, and MAE represents the mean annual evaporation. The red line represents the tree level, the green line is the shrub level, and the blue line is the herb level. R

^{2}represents how well the model fits the variables studied and the p-value represents the significance level.

**Figure 6.**The general linear analysis of soil factors with different life forms of LN. (

**a**) General linear relationship between the soil N and LN of plants in the trees, shrubs, and herbs. (

**b**) General linear relationship between the soil P and LN of plants in the trees, shrubs, and herbs. (

**c**) General linear relationship between the soil pH and LN of plants in the trees, shrubs, and herbs. The red line represents the tree level, the green line is the shrub level, and the blue line is the herb level. R

^{2}represents how well the model fits the variables studied and p-value represents the level of significance.

**Figure 7.**The general linear analysis of soil factors with different life forms of LP. (

**a**) General linear relationship between the soil N and LP of plants in the trees, shrubs, and herbs (

**b**) General linear relationship between the soil P and LP of plants in the trees, shrubs, and herbs. (

**c**) General linear relationship between the soil pH and LP of plants in the trees, shrubs, and herbs. The red line represents the tree level, the green line is the shrub level, and the blue line is the herb level. R

^{2}represents how well the model fits the variables studied and the p-value represents the level of significance.

**Figure 8.**The general linear analysis of soil factors with different life forms of N/P. (

**a**) General linear relationship between the soil N and N/P of plants in the trees, shrubs, and herbs (

**b**) General linear relationship between the soil P and N/P of plants in the trees, shrubs, and herbs. (

**c**) General linear relationship between the soil pH and N/P of plants in the trees, shrubs, and herbs. The red line represents the tree level, the green line is the shrub level, and the blue line is the herb level. R

^{2}represents how well the model fits the variables studied and the p-value represents the level of significance.

**Figure 9.**The NMDS ranking of climatic and soil factors with different life forms of LN. Value of de represents the deviation explained by the corresponding model. (

**a**) NMDS ranking of soil factors with tree levels LN; (

**b**) NMDS ranking of soil factors with shrub levels LN; (

**c**) NMDS ranking of soil factors with herb levels LN; (

**d**) NMDS ranking of climatic factors with tree levels LN; (

**e**) NMDS ranking of climatic factors with shrub levels LN; (

**f**) NMDS ranking of climate factors and herb levels LN; (

**g**) NMDS ranking of the sum of soil factors and climate factors and tree levels LN; (

**h**) NMDS ranking of the sum of soil factors and climate factors and shrub levels LN; (

**i**) NMDS ranking of the sum of soil factors and climate factors and herb levels LN. Trait stacking indicates that abiotic factors, indicated by points on the NMD, are associated with higher or lower trait values, consistent with a colored trait gradient. Note that if the relationship between the LN and abiotic factors is linear, the gradient splines will be parallel. Nonlinear relationships between LN and abiotic factors are represented by curve splines.

**Figure 10.**The NMDS ranking of climatic and soil factors with different life forms of LP. Value of de represents the deviation explained by the corresponding model. (

**a**) NMDS ranking of soil factors with tree levels LP; (

**b**) NMDS ranking of soil factors with shrub levels LP; (

**c**) NMDS ranking of soil factors with herb levels LP; (

**d**) NMDS ranking of climatic factors with tree levels LP; (

**e**) NMDS ranking of climatic factors with shrub levels LP; (

**f**) NMDS ranking of climate factors and herb levels LP; (

**g**) NMDS ranking of the sum of soil factors and climate factors and tree levels LP; (

**h**): NMDS ranking of the sum of soil factors and climate factors and shrub levels LP; (

**i**): NMDS ranking of the sum of soil factors and climate factors and herb levels LP. Trait stacking indicates that abiotic factors, indicated by points on the NMD, are associated with higher or lower trait values, consistent with a colored trait gradient. Note that if the relationship between LP and abiotic factors is linear, the gradient splines will be parallel. Nonlinear relationships between LP and abiotic factors are represented by curve splines.

**Figure 11.**The NMDS ranking of climatic and soil factors with different life forms of N/P. Value of de represents the deviation explained by the corresponding model. (

**a**) NMDS ranking of soil factors with tree levels N/P; (

**b**) NMDS ranking of soil factors with shrub levels N/P; (

**c**) NMDS ranking of soil factors with herb levels N/P; (

**d**) NMDS ranking of climatic factors with tree levels N/P; (

**e**) NMDS ranking of climatic factors with shrub levels N/P; (

**f**) NMDS ranking of climate factors and herb levels N/P; (

**g**) NMDS ranking of the sum of soil factors and climate factors and tree levels N/P; (

**h**) NMDS ranking of the sum of soil factors and climate factors and shrub levels N/P; (

**i**) NMDS ranking of the sum of soil factors and climate factors and herb levels N/P. Trait stacking indicates that abiotic factors, indicated by points on the NMD, are associated with higher or lower trait values, consistent with a colored trait gradient. Note that if the relationship between N/P and abiotic factors is linear, the gradient splines will be parallel. Nonlinear relationships between N/P and abiotic factors are represented by curve splines.

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**MDPI and ACS Style**

Wang, X.; Wang, J.; Zhang, L.; Lv, C.; Liu, L.; Zhao, H.; Gao, J.
Climatic Factors Determine the Distribution Patterns of Leaf Nutrient Traits at Large Scales. *Plants* **2022**, *11*, 2171.
https://doi.org/10.3390/plants11162171

**AMA Style**

Wang X, Wang J, Zhang L, Lv C, Liu L, Zhao H, Gao J.
Climatic Factors Determine the Distribution Patterns of Leaf Nutrient Traits at Large Scales. *Plants*. 2022; 11(16):2171.
https://doi.org/10.3390/plants11162171

**Chicago/Turabian Style**

Wang, Xianxian, Jiangfeng Wang, Liuyang Zhang, Chengyu Lv, Longlong Liu, Huixin Zhao, and Jie Gao.
2022. "Climatic Factors Determine the Distribution Patterns of Leaf Nutrient Traits at Large Scales" *Plants* 11, no. 16: 2171.
https://doi.org/10.3390/plants11162171