Ecological differentiation and changes in historical distribution of Corylus heterophylla species complex since the last interglacial
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摘要:目的 探讨末次间冰期以来平榛物种复合体（平榛、川榛、滇榛）的历史分布变迁，了解其地理分布对环境演变的响应机制，旨在为厘清3个物种的进化关系提供借鉴意义，也为资源保护和开发提供科学依据。方法 基于平榛物种复合体3个近缘种的分布数据及相应的筛选后的环境变量，利用MaxEnt模型及ArcGIS软件对平榛物种复合体在末次间冰期、末次盛冰期、全新世中期和现代的潜在分布区进行模拟，以探究其历史分布变迁，并划分适生区等级；将3个近缘种在不同时期的分布区域进行叠加，推测其避难所，并采用榛孢粉数据进行验证；最后，利用主成分分析法探究3个近缘种的生态位分化情况，并对主要环境影响因子进行综合评价。结果 从整体来看，平榛和川榛的前2位主要环境因子贡献度相差不大，且滇榛前2位主要环境因子均与水分有关，相对于温度，水分对3种树种的分布有着更重要的影响。从适生区面积来看，平榛的适生区面积在冰期呈收缩趋势，进入全新世中期后面积又有所扩张，川榛和滇榛的适生区面积却与之相反；适生区区域重叠后得到的3个近缘种的避难所与孢粉信息较吻合。结论 本研究较为准确地模拟了平榛物种复合体在4个时期的适生区分布变迁，探究了其生态位分化情况，并推测了其生物避难所，可为未来气候变化背景下中国境内榛资源保护和管理开发提供一定的科学依据。Abstract:Objective This paper aims to discuss the historical distribution changes of Corylus heterophylla complex (Corylus heterophylla, Corylus kweichowensis, Corylus yunnanensis) since the last interglacial, and understand the response mechanism of its geographical distribution to environmental evolution, so as to provide reference for clarifying the evolutionary relationship of three species, and also provide scientific basis for resource protection and development.Method Based on the distribution data of the three species of C. heterophylla complex and the corresponding environmental variables, MaxEnt model and ArcGIS software were used to simulate their potential distribution in the last interglacial, last glacial maximum, mid-Holocene and modern times to explore its historical distribution changes; then, the distribution areas of three related species in different periods were superimposed to speculate their ice age refuge, which was verified by hazel pollen data. Finally, the niche differentiation among the three related species was detected by principal component analysis, and the main environmental impact factors were comprehensively evaluated.Result On the whole, the contribution of the first two environmental factors affecting the distribution of C. heterophylla and C. kweichowensis was similar, while the first two main environmental factors of C. yunnanensis were related to water. Compared with temperature, water had a more important impact on the distribution of the three tree species. In terms of the suitable areas, C. heterophylla of northern China experienced glacial contraction and interglacial expansion during the quaternary, whereas C. kweichowensis and C. yunnanensis of southern China presented population expansion even during the last glacial maximum. The refuges of the three related species obtained after overlapping of the suitable areas were in good agreement with the palynological results.Conclusion This study accurately simulates the distribution changes of C. heterophylla complex in four historical periods, explores its ecological differentiation, and speculates on its biological refuge, which provides some scientific basis for the conservation and management of the resources in China under the background of future climate change.
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图 2 基于304个物种分布点12个气候变量的主成分分析（PCA）
Figure 2. PCA of 12 climate variables based on 304 species distribution points
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图 3 平榛物种复合体4个时期分布区预测
底图审图号：GS(2019)1822。下同。Base drawing review No: GS(2019)1822. The same below.
Figure 3. Prediction of the distribution area of C. heterophylla species complex in 4 periods
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图 4 平榛物种复合体避难所预测及相近孢粉数据点
Figure 4. Prediction of C. heterophylla species complex shelter and similar sporopollen data points
表 1 平榛物种复合体筛选所用气候变量
Table 1 Filter climate variable used by C. heterophylla complex
树种 Tree species 变量代码 Variable code 气候变量 Climate variable 平榛 C. heterophylla bio2 昼夜温差月均值 Monthly mean diurnal difference of temperature bio3 等温性 Isothermality bio4 温度季节变化 Seasonal change in temperature bio10 最暖季度平均温度 Mean temperature of the warmest quarter bio15 降水量变异系数 Coefficient of variation of precipitation bio16 最湿季度雨量 Precipitation of the wettest quarter bio17 最干季度雨量 Precipitation of the driest season 川榛 C. kweichowensis bio3 等温性 Isothermality bio4 温度季节变化 Seasonal change in temperature bio5 最热月份最高温 Max. temperature of the warmest month bio6 最冷月份最低温 Min. temperature of the coldest month bio12 年平均雨量 Annual mean precipitation bio15 降水量变异系数 Coefficient of variation of precipitation 滇榛 C. yunnanensis bio4 温度季节变化 Seasonal change in temperature bio6 最冷月份最低温 Min. temperature of the coldest month bio7 年温度变化范围 Temperature annual range bio12 年平均雨量 Annual mean precipitation bio19 最冷季度雨量 Precipitation of the coldest season 
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表 2 模型AUC预测精度
Table 2 Modeling prediction precision of AUC
树种
Tree species平均AUC值
Average AUC value气候模型 Climate model 末次间冰期
Last interglacial (LIG)末次盛冰期
Last glacial maximum (LGM)全新世中期
Mid holocene (MH)现在气候
Current climate平榛 C. heterophylla 训练集 Training data 0.990 0.992 0.991 0.991 测试集 Testing data 0.983 0.988 0.987 0.988 川榛 C. kweichowensis 训练集 Training data 0.990 0.990 0.990 0.989 测试集 Testing data 0.989 0.991 0.989 0.989 滇榛 C. yunnanensis 训练集 Training data 0.996 0.996 0.995 0.995 测试集 Testing data 0.997 0.997 0.997 0.996
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表 3 主要气候因子贡献度
Table 3 Contribution of major climate factors
树种
Tree species变量代码
Variable code气候变量
Climate variable置换重要值
Substitution of important value累计贡献率
Cumulative contribution rate/%平榛 C. heterophylla bio4 温度季节变化 Seasonal change of temperature 54.2 31.1 bio16 最湿季度雨量 Precipitation of the wettest season 31.3 30.0 bio15 降水量变异系数 Coefficient of variation of precipitation 0.8 26.2 川榛 C. kweichowensis bio4 温度季节变化 Seasonal change of temperature 9.0 26.7 bio12 年平均雨量 Annual mean precipitation 9.5 26.3 bio15 降水量变异系数 Coefficient of variation of precipitation 24.3 22.8 滇榛 C. yunnanensis bio12 年平均雨量 Annual mean precipitation 13.1 36.9 bio19 最冷季度雨量 Precipitation of the coldest season 24.7 23.8 bio4 温度季节变化 Seasonal change of temperature 11.2 17.6
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表 4 气候变量指标PCA特征值及方差
Table 4 PCA eigenvalues and variances of climate variable indicators
主成分
Principal component特征值
EigenvalueRV/% RCV/% 1 6.482 44 54.020 31 54.020 31 2 2.695 33 22.461 10 76.481 40 3 1.237 97 10.316 44 86.797 85 4 0.765 71 6.380 95 93.178 79 5 0.470 02 3.916 83 97.095 63 6 0.178 25 1.485 42 98.581 05 7 0.131 52 1.095 97 99.677 02 8 0.019 66 0.163 86 99.840 88 9 0.009 32 0.077 68 99.918 57 10 0.005 35 0.044 57 99.963 14 11 0.003 74 0.031 20 99.994 34 12 67919.7 0.005 66 100.000 00 注：RV. 方差百分比；RCV. 累计贡献度。Notes: RV, percentage of variance; RCV, cumulative contribution.
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表 5 平榛物种复合体分布区内出现的孢粉点
Table 5 Sporopollen points appearing in the distribution area of C. heterophylla species complex
序号
No.采集地
Collecting site经纬度
Coordinate地层时期
Chronologic period文献
Reference1 北京市顺义区 Shunyi District, Beijing 40°09′00″N，116°32′00″E LIG, LGM, MH [19] 2 北京市大兴区 Daxing District, Beijing 39°43′00″N，116°19′00″E LIG, LGM, MH [19] 3 陕西省富县 Fuxian County, Shaanxi Province 36°02′00″N，109°17′00″E LIG, LGM [20] 4 吉林省敦化市 Dunhua City, Jilin Province 43°22′00″N，128°13′00″E MH [21] 5 吉林省靖宇县 Jingyu County, Jilin Province 42°09′00″N，126°23′00″E MH [22] 6 云南省鹤庆县 Heqing County, Yunnan Province 26°33′00″N，100°10′00″E LIG, LGM [23] 7 湖南省城步苗族自治区
Chengbu Miao Autonomous Region, Hunan Province26°23′00″N，110°19′00″E MH [24] 8 贵州省荔波县 Libo County, Guizhou Province 25°24′00″N，107°52′00″E MH [25] 9 贵州省镇宁布依族苗族自治县
Buyi Miao Autonomous County, Zhenning, Guizhou Province26°02′00″N，105°50′00″E MH [26] 10 湖北省神农架林区 Shennongjia Forest District, Hubei Province 31°29′00″N，109°59′00″E LGM, MH [27] 11 浙江省杭州市 Hangzhou City, Zhejiang Province 30°08′00″N，120°13′00″E MH [28] 12 甘肃省渭源县 Weiyuan County, Gansu Province 35°08′00″N，104°12′00″E LGM, MH [29] 13 陕西省武功县 Wugong County, Shaanxi Province 34°15′00″N，108°12′00″E LIG [30] 14 陕西省蓝田县 Lantian County, Shaanxi Province 34°09′00″N，109°19′00″E LIG [30] 15 陕西省渭南阳郭 Yangguo, Weinan, Shaanxi Province 34°21′00″N，109°31′00″E LGM [31] 16 四川省布拖县 Butuo County, Sichuan Province 27°42′00″N，102°49′00″E LGM, MH [32] 17 陕西省太白县 Taibai County, Shaanxi Province 34°03′00″N，107°18′00″E LGM [33]
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表 6 不同时期平榛复合体在中国的适生区预测
Table 6 Prediction of suitable area of C. heterophylla complex in China in different periods
104 km2 树种
Tree species时间段
Period总适生区
Total suitable
region低度适生区
Lowly suitable
region一般适生区
Generally suitable
region中度适生区
Moderately suitable
region高度适生区
Highly suitable
region平榛 C. heterophylla LIG 161.351 7 78.066 5 41.404 0 22.911 3 18.969 8 LGM 150.652 8 76.902 8 28.239 6 28.701 4 16.809 0 MH 139.855 9 68.651 0 34.099 0 18.626 7 18.479 2 现代 Current 133.562 5 57.092 0 34.467 0 26.269 1 15.734 4 川榛 C. kweichowensis LIG 169.376 0 91.397 2 36.447 3 31.336 3 10.195 1 LGM 176.685 8 93.017 4 38.222 2 25.326 4 20.119 8 MH 163.218 8 77.012 2 36.149 3 27.533 0 22.524 3 现代 Current 170.850 7 62.338 5 50.588 5 33.689 2 24.234 4 滇榛 C. yunnanensis LIG 54.332 2 27.162 0 16.434 9 7.256 7 3.478 5 LGM 97.474 0 60.684 0 19.000 0 11.734 4 6.055 6 MH 71.720 5 40.036 5 18.156 3 8.614 6 4.913 2 现代 Current 76.942 7 46.048 6 13.107 6 13.430 6 4.355 9
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