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For modelling purposes, we selected 20 native/naturalized tree species ethnobotanically important as fodder resources and commonly used in afforestation or in social forestry in China. A review of the literature (ESM 1) revealed that most of these species were multipurpose, fast growing and beneficial for the soil through various combinations of N-fixation, C-sequestration, soil stabilization and erosion prevention (Table 1). Many of the selected trees were already used in agroforestry systems as shelterbelts, windbreaks, or for improving crop and livestock production through alley cropping. In addition to adding fodder value for livestock, selected trees were or could be used as: edible (12), timber (12), medicinal (10), fiber (10), bioenergy feedstock (11) and industrial value (5) (Table 1).
Table 1. Characteristic of selected fodder trees, ethnobotanical notes, and agroforestry uses.
Species Growth rate Fodder value Soil improvement Potential economic uses Agroforestry Ailanthus altissima Fast High especially for goat, good for silkworm C-sequestration, soil stabilization Medicinal, timber, fuelwood Shelterbelt, potential for cultivation in heavily polluted areas and drought tolerant; known to have allelopathic effect, and therefore, proper management is necessary Amorpha fruticosa Fast High, bee forage N-fixation, Erosion control Medicinal, edible, industrial uses Shelterbelt, tolerates poor sandy soils, dry soils, limey soils, acidic soils Arundo donax Fast High C-sequestration, soil stabilization Bioenergy feedstock, medicinal, thatching Shelterbelt, windbreak, ability to grow in different soil types and climatic conditions Boehmeria clidemioides Fast High Remediate soils with heavy metals contamination Fiber, medicinal Local ethnobotanical value, planted in gullies Boehmeria nivea Fast High Prevent soil erosion Important fiber crop in China, medicinal, edible Planted in gullies Broussonetia papyrifera Fast High Increase phosphorus and nitrogen and improve soil moisture Food, paper making, bioenergy feedstock, fiber, medicinal, timber Shelterbelt and windbreak, economic fallow crop, leading to increased crop production Castanea mollissima Medium High Increase organic matter, nitrogen, phosphorus, and potassium content Edible, bioenergy feedstock, produce utilizable timber every 10 year Alley cropping, silviculture practices, good results from Castanea-tea intercropping Cyclobalanopsis glauca Medium High C-sequestration, improve soil nitrogen Fuelwood, bioenergy feedstock, timber Branch and twigs are good material for mushroom culture Debregeasia orientalis Fast High Improve metal contaminated soil Edible, high-quality fiber Local ethnobotanical value Elaeagnus angustifolia Fast Medium, good bee forage N-fixation, Erosion control or dune stabilization Edible, industrial value, bioenergy feedstock, timber Shelterbelts, windbreaks or protective plantings Ficus heteromorpha Fast Medium Stabilize soil and increase fertility of soil Medicinal, edible, paper making, pig feed Shelterbelts, windbreaks Leucaena leucocephala Fast High, good bee forage N-fixation, C-sequestration Fiber, edible, timber Very good for a maize crop, alley cropping systems Machilus gamblei Fast High, used for Muga silkworm (Antherea assamensis) in NE India N-fixation, C-sequestration Edible, fiber, medicinal, timber, potential for bioenergy feedstock Local ethnobotanical value Morus alba Fast High, good for silkworm Erosion control Edible, industrial value, bioenergy feedstock Shade and shelter, windbreak Populus adenopoda Fast High Increase in soil organic carbon, soil stabilization Timber, fiber, bioenergy feedstock Shade and shelter, windbreak Populus davidiana Fast High Increase in soil organic carbon, soil stabilization Timber, bioenergy feedstock Shade and shelter, windbreak Populus tomentosa Fast High Increase in soil organic carbon, soil stabilization Timber, fiber, bioenergy feedstock Shade and shelter, windbreak Salix babylonica Fast High, bee forage Erosion control Medicinal, fiber, light timber, bioenergy feedstock Shade and shelter, windbreak Saurauia thyrsiflora Fast High Erosion control Edible, medicinal Local ethnobotanical value, high milk production in livestock Ulmus pumila Fast High Erosion control, stabilizing sand dunes Fiber, medicinal, edible, timber Shelterbelt, windbreak, enhance crop production Source: literature listed in ESM 1 Climatic suitability of trees and crops
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The stepwise elimination of bioclimatic variables with VIF values greater than 10 resulted in a set of ten least correlated (Pearson correlation value < 0.8) bioclimatic variables: bio2, bio3, bio8, bio13, bio14, bio15, bio21, bio22, bio25 and bio31. These variables successfully produced a distribution model for each of the selected tree species. Consensus maps showing the results from ensemble model for each fodder tree species are given in Figure 2. The models correctly predicted most of the test locations in all cases. Sub-models for all selected species weight between 0.05 and 0.29 (Table 2). The ensemble models had final AUC ranges from 0.82 to 0.98 for different species, while kappa values ranged from 0.74 to 0.97 (Table 2).
Figure 2. Consensus mapping based on ensemble modelling. The bioclimatic suitability for each of the selected species, with the observed point distributions overlain. ‘a’ to ‘t’ are species codes for the fodder tree species as listed in Table 2.
Table 2. Final weights of each sub-models for ensemble forecasting, AUC, kappa and threshold for ensemble models
GLM MAXENT MAXLIKE RF RPART ENSEMBLE Spp code Spp Weight Weight Weight Weight Weight AUC Kappa maxTPR+TNR a Ailanthus altissima 0.22 0.24 0.22 0.23 0.09 0.84 0.81 0.59 b Amorpha fruticosa 0.25 0.26 0.22 0.26 0 0.82 0.74 0.57 c Arundo donax 0.24 0.25 0.17 0.25 0.09 0.93 0.84 0.6 d Boehmeria clidemioides 0.22 0.22 0.15 0.22 0.19 0.92 0.85 0.48 e Boehmeria nivea 0.2 0.22 0.21 0.22 0.16 0.92 0.89 0.53 f Broussonetia papyrifera 0.23 0.24 0.1 0.24 0.19 0.92 0.82 0.56 g Castanea mollissima 0.21 0.21 0.2 0.21 0.16 0.88 0.83 0.65 h Cyclobalanopsis glauca 0.2 0.21 0.2 0.21 0.18 0.92 0.86 0.51 i Debregeasia orientalis 0.19 0.21 0.21 0.21 0.18 0.95 0.94 0.46 j Elaeagnus angustifolia 0.21 0.22 0.15 0.23 0.18 0.89 0.85 0.61 k Ficus heteromorpha 0.21 0.21 0.21 0.21 0.16 0.91 0.81 0.55 l Leucaena leucocephala 0.24 0.25 0.17 0.26 0.08 0.94 0.93 0.68 m Machilus gamblei 0.21 0.24 0.23 0.23 0.08 0.97 0.97 0.75 n Morus alba 0.25 0.25 0.25 0.24 0 0.84 0.85 0.64 o Populus adenopoda 0.29 0.29 0.05 0.29 0.09 0.89 0.9 0.54 p Populus davidiana 0.22 0.22 0.21 0.23 0.12 0.85 0.89 0.44 q Populus tomentosa 0.21 0.24 0.21 0.26 0.09 0.89 0.81 0.6 r Salix babylonica 0.18 0.26 0.24 0.27 0.06 0.83 0.74 0.55 s Saurauia thyrsiflora 0.19 0.21 0.2 0.21 0.18 0.98 0.97 0.76 t Ulmus pumila 0.22 0.22 0.2 0.22 0.13 0.84 0.86 0.51 NB: sub-model with ‘0’ indicate that particular sub-model was calibrated but not used in the ensemble model. Bioclimatic suitability of 20 fodder tree species within China as estimated by the ensemble modelling is presented in Figure 2 (also listed in ESM 2). Species were classified based on the area of bioclimatic suitability and magnitude of suitability. Suitability of fodder trees was well represented across each of the seven crop growing areas (Fig. 2 and Fig. 3) in China. More fodder species found bioclimatic suitability in areas where grain crops, crop/pasture rotations, agro-silvopastoral systems and ponds systems dominated. Few of the selected fodder tree species showed bioclimatic suitability in areas dominated by rangeland systems.
Figure 3. Climatic suitability index of important crops in China; Millet include both pearl and foxtail millet.
The climate suitability index of selected crops was found to be relatively low for northern and western China where low yield was reported. Higher climate suitability index was found for all the crops in southern and eastern parts of China comprised of mostly overlapping farmlands and adjoining areas (Fig. 3).
Integration possibilities
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The fuzzy logic model output identified areas where fodder trees could be integrated with livestock and crops. According to the model, rice-based integration could be suitable in the southern, southeast and eastern parts of China. Maize, soybean and sorghum could be integrated in the southern, eastern and central parts, whereas millet, wheat and sorghum could be integrated in central and northern parts. Distinct spatial data were available for different livestock species, whereas such data were lacking for individual crops, meaning that crops could not be treated individually like livestock and fodder tree species. The potential distribution of each fodder tree was overlain with the distributions of different livestock species and crops to produce fuzzy maps (Fig. 4; for individual livestock detail maps documented in ESM 3). The range of the fuzzy maps lies between ‘0’ and ‘1’. When membership function values of all the three layers were 0 (minimum), the suitability for integration was minimum, and when the membership function values were 1 (maximum) the possibility for integration was at its greatest. Our model revealed that most of the fodder tree species were suitable for integration toward the eastern and southern parts of China. Few species found agro-ecological suitability in northern China, and few species could be incorporated into integrated systems in western China (Fig. 4). The most suitable areas fell in the humid and sub-humid regions, with few options for integration in the more arid regions.
Figure 4. Major areas suitable for integration of selected fodder trees with crops and livestock. ‘a’ to ‘t’ are species codes for the fodder species as listed in Table 2.
Characteristic of modelled trees
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We considered the whole of China for the modelling work. Agriculture is a vital sector in China, providing livelihoods to several hundred million people. Rice, corn, wheat, sorghum and soybeans are among the major crop produced in China. Mixed crop-livestock production systems are commonly practiced, and five broad production systems are recognized[8]. These systems are separated based on agro-ecology, dividing the country into rangeland, grain crops, crop/pasture rotations, agro-silvopastoral systems and pond systems, overlain with the seven important crops mentioned above (Fig. 5).
Figure 5. Map of eco-regions, cropland and the crop-livestock production systems in China: 1. systems based on rangeland; 2. systems based on grain crops; 3. systems based on crop/pasture rotations; 4. agro-silvopastoral systems; and 5. systems based on ponds (adapted from Hou et al.[8] and Broxton et al.[52])
Data
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In this paper, we included 20 tree species. These were selected based on (i) existing plantation practice and farmers' preferences; (ii) well-perceived potential of the species to address economic and ecological benefits (e.g., livelihood source, soil fertility, shade tree and combinations thereof); and (iii) availability of ground data from various ethnobotanical and agroforestry projects in which the authors were involved as well as a review of the relevant literature (e.g., Fang et al.[53]). The twenty fodder tree species were represented by a total 3,016 geo-coordinated points which were consigned to a 2 km grid for the analysis. Cropland data were extracted using the Land Use Land Cover (LULC) 2010 map of China. Crops (Table 3) were modelled based on temperature and precipitation relations. Livestock data were obtained from Robinson et al.[54] and confirmed with annual statistical information from the Chinese government. Livestock used in this study includes cattle, buffaloes, goats and sheep. We used bio-climatic variables downloaded from the CliMond archive[55] as inputs for the modelling of the 20 fodder tree species. Detailed information on these bio-climatic variables, comprising variables of temperature, precipitation, radiation and moisture indexes are listed in ESM 4.
Table 3. Optimum and absolute range of temperature and precipitation for selected crops.
Absolute Optimum Crop Tmax Tmin Pmax Pmin Tmax Tmin Pmax Pmin Oryza sativa Rice 36 10 4 000 1 000 30 20 2 000 1 500 Sorghum bicolor Sorghum 40 8 3 000 300 35 27 1 000 500 Zea mays Maize 47 10 1 800 400 33 18 1 200 600 Hordeum vulgare Barley 40 2 2 000 200 20 15 1 000 500 Pennisetum glaucum Pearl millet 40 12 1 700 200 35 25 900 400 Setaria italica Foxtail millet 35 5 4 000 300 26 16 700 500 Glycine max Soybean 38 10 1 800 450 33 20 1 500 600 Triticum aestivum Wheat 27 5 1 600 300 23 15 900 750 Tmax − Maximum temperature, Tmin − Minimum temperature in °C; Pmax − Maximum precipitation, Pmin − Minimum precipitation in mm Tree and crop distribution modelling
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BiodiversityR package (Ver. 2.8−2) and dismo package in R were used to prepare bioclimatic suitability maps for the 20 fodder trees, important crops and other analysis. Five different algorithms − Maxent, Maxlike, Random Forests (RF), Generalized Linear Models (GLM) and Recursive Partitioning and Regression Trees (RPart) − were used in the tree modelling. Following Hijmans[56], we used 4-fold cross-validation, where tree occurrence and background data were partitioned into 75% calibration and 25% evaluation observations. The consensus mapping technique is based on an ensemble of several niche-modeling algorithms (sub-models). The results using each model were treated as individual sub-models (Pmod), with weights assigned based on their performance. An ensemble model (Pensemble) was then calculated using the following formula[57,58]:
$ {P}_{ensemble}=\frac{\sum ({w}_{mod}{P}_{mod})}{\sum \left({w}_{mod}\right)} $ (1) where, wmod = weighted averages of sub-models (Pmod).
Bioclimatic variables were selectively removed based on variance inflation factor (VIF)[59] calculations where VIF > 10 were eliminated (ESM 5) to provide a minimum set of the least correlated bioclimatic variables.
An EcoCrop model was used to identify areas suitable for selected seven important crops in China. The EcoCrop is a simple mechanistic model that use expert-based temperature and rainfall ranges[60]. The FAO-EcoCrop database (http://ecocrop.fao.org/) provided such ranges for crops and hence used in this work to determine the climatic niche of crops in China and then produces a suitability score. The model needs absolute range (at which the crop can grow) and the optimum range (at which the crop grows best) of temperature and precipitation. Table 3 presents range of absolute and optimum temperature and precipitation for selected crops. The model determines suitability index based on the conditions over the growing season at a particular place using a gridded data of temperature and rainfall. The suitability index ranges from zero (not suitable) to 100 (highly suitable). The model performs two different calculations separately, one for rainfall and the other for temperatures, and then calculates the interaction by multiplying them.
Model evaluation
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We evaluated the ability of sub-models to cope with spatial autocorrelation by calculating calibrated Area Under the Receiver Operator Curve (cAUC) values and comparing these with a geographical null mode[59]. Spatial sorting bias[56] was removed and carried out through several rounds of calibration and evaluation of all models (including the geographical null model), each time using three partitions for model calibration and one partition for model evaluation. Elimination of spatial sorting bias in testing data in model calibrations produced cAUC values of the different algorithms between 0.6 to 0.85 for different tree species. These values were significantly different from the null model (0.49 and 0.501 for the null model, which is equivalent to a random draw[56]; Mann-Whitney tests, p < 0.05 in all cases). Weights calculated for the sub-models were used to determine the appropriate weights (ranging between 0 and 1) for the ensemble model.
Kappa and AUC values were calculated for each of the ensemble models. The ensemble output consists of a consensus map that represents the agreement between sub-models. All pixels in the consensus map output were classified according to the cut-off point, based on a threshold defined by maximizing the sum of the true presence and true absence rates (maxTPR+TNR). A score above this threshold represents the suitable climatic space for the species in question[57]. All pixels with suitability scores above the cut-off point were included in the final bioclimatic suitability map for each species.
Predicting potential zones for mixed plantations
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A fuzzy logic model was employed to identify areas suitable for tree-crop-livestock integration. Classified raster layers of fodder tree and crop distributions were overlain with raster layers of ruminant livestock and cropland in China (Fig. 5). Fuzzy overlay analysis is based on set theory in which a set generally corresponds to a class. Fuzzy overlay analysis reclassifies or transforms data values to a common scale, but the transformed values represent the probability of belonging to a specified class. Fuzzy logic system can facilitate complex approaches, such as the incorporation of biotic interaction in the modelling[28]. The combining step in fuzzy overlay analysis quantifies each location’s probability of belonging to specified sets from various input rasters[24,29]. The equation using fuzzy Gaussian function is:
$ \mu \left(x\right)={e}^{-{f}_{1}×{\left(x-{f}_{2}\right)}^{2}} $ (2) where, the inputs to the equation f1 and f2 are the spread and the midpoint, respectively. The midpoint is a user-defined value with a fuzzy membership of 1. The default is the midpoint of the range of values of the input raster. Spread defines the membership of the Gaussian function. It generally ranges from 0.01 to 1. Increasing the spread causes the fuzzy membership curve steeper. Fuzzy overlay analysis quantifies the possibilities of each cell or location to a specified set based on membership value[29].
Study area
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The research was financially supported by key research project “Agroforestry Systems for restoration and bio-industry technology development (grant no: 2017YFC0505101)”, by the Agriculture Science and Technology Innovation Program (ASTIP-IAS07; CAAS-XTCX2016011-01), Research Program of the State Key Laboratory of Animal Nutrition (2004DA125184G1103), Bureau of International Cooperation Chinese Academy of Sciences (151853KYSB20160032) and CGIAR Research Program on Climate Change (FTA-FP5). We acknowledge support from Key Laboratory of Economic Plants and Biotechnology Kunming Institute of Botany, Chinese Academy of Sciences Kunming China and Prof. Yuhua Wang for providing information and data on fodder trees.
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