林业科学 ›› 2020, Vol. 56 ›› Issue (4): 1-11.doi: 10.11707/j.1001-7488.20200401
张晓玮1,2,王婧如2,王明浩2,杨毅2,赵长明2
收稿日期:
2018-05-07
出版日期:
2020-04-25
发布日期:
2020-05-26
基金资助:
Xiaowei Zhang1,2,Jingru Wang2,Minghao Wang2,Yi Yang2,Changming Zhao2
Received:
2018-05-07
Online:
2020-04-25
Published:
2020-05-26
摘要:
目的: 以我国各地理阶梯间山地生长的云杉属12个树种为对象,探讨限制云杉地理分布和驱动各树种沿经纬度分布差异的关键气候因子,以阐明影响我国云杉属各树种地理分布格局形成的主导气候因子。方法: 在长期野外考察积累和查阅中国数字植物标本馆标本信息的基础上,于云杉属12个树种的地理分布范围内选取389个样点,并利用地理信息系统技术(ArcGIS)获取各样点的主要气候数据。通过方差分析、变异系数比较确定限制云杉树种地理分布范围的气候因子,基于线性回归分析、蒙特卡洛检验和冗余分析(RDA)等方法量化各气候因子对云杉树种地理分布差异的贡献大小。结果: 年均气温、月均昼夜气温差、最暖月最高气温、最冷月最低气温、年均气温变幅、年均降水量、最暖季降水量、最冷季降水量和干旱指数在12个树种分布区间存在显著差异;月均昼夜气温差和最冷季降水量的方差主要来源于树种内,其余7个气候因子的方差主要来源于树种间;变异系数分析表明,无论是云杉属整体分布区还是各树种分布区的月均昼夜气温差和最暖月最高气温变异最低,其变异系数均小于20%;经、纬度与各气候因子的线性回归分析表明,除经、纬度与月均昼夜气温差、经度与最冷季降水量不存在显著相关(P>0.05)外,经、纬度与其余各气候因子均显著相关(P < 0.05),其中,与年均气温变幅相关性最高,其次为最冷月最低气温;蒙特卡洛检验进一步表明,年均气温变幅和最冷月最低气温对我国云杉属各树种沿经纬度分布差异的解释率分别达到84%和66.8%;RDA分析显示,第1主分轴主要反映热量条件,其中重要的气候因子分别为年均气温变幅、最暖月最高气温和最冷月最低气温,其载荷值分别为0.93、-0.83和0.64,第2主分轴主要反映以年均降水量、干旱指数和最暖季降水量为主的水分情况解释的信息量,且前2主分轴对树种和树种-环境关系的累积解释率分别高达89.2%和100%。结论: 限制我国云杉属树种地理分布的主要气候因子为月均昼夜气温差和最暖月最高气温,而导致各树种沿经纬度分布存在明显地理差异的主要驱动因子为年均气温变幅和最冷月最低气温。本研究进一步证实热量是影响云杉属植物等寒温性植物地理分布格局形成的主要原因和差异来源,降水量在一定程度上只起到次要作用。
中图分类号:
张晓玮,王婧如,王明浩,杨毅,赵长明. 中国云杉属树种地理分布格局的主导气候因子[J]. 林业科学, 2020, 56(4): 1-11.
Xiaowei Zhang,Jingru Wang,Minghao Wang,Yi Yang,Changming Zhao. Dominant Climatic Factors Influencing the Geographical Distribution Pattern of Picea in China[J]. Scientia Silvae Sinicae, 2020, 56(4): 1-11.
表1
所选云杉属各树种的分布地区、经纬度范围及样点数"
树种Tree species | 分布地区Distribution region | 经度 Latitude (E)/(°) | 纬度 Longitude (N)/(°) | 样点数 Sample number |
鱼鳞云杉Picea jezoensis | 黑龙江Heilongjiang、吉林Jilin、内蒙古Inner Mongolia | 123.7-134.0 | 41.2-48.2 | 20 |
红皮云杉P. koraiensis | 黑龙江Heilongjiang、吉林Jilin、辽宁Liaoning | 126.5-130.5 | 41.4-47.0 | 20 |
白杄P. meyeri | 山西Shanxi、河北Hebei、内蒙古中部Central Inner Mongolia | 111.1-110.3 | 37.3-44.0 | 23 |
青杄P. wilsonii | 内蒙古中部Central Inner Mongolia、河北Hebei、山西Shanxi、陕西南部Southern Shaanxi、湖北西部Western Hubei、甘肃中南部South central Gansu | 101.6-116.8 | 33.7-40. 8 | 53 |
青海云杉P. crassifolia | 青海东北部Northeast Qinghai、甘肃Gansu、宁夏Ningxia、内蒙古中部Central Inner Mongolia | 100.9-104.3 | 33.6-37.8 | 34 |
云杉P. asperata | 陕西西南部Southwest Shaanxi、甘肃东部和南部Eastern and Southern Gansu、四川Sichuan | 101.1-104.4 | 30.3-37.8 | 33 |
丽江云杉 P. likiangensis var. likiangensis | 云南西北部Northwest Yunnan、四川西南部Southwest Sichuan、西藏东南部Southeast Tibet | 98.8-101.9 | 26.8-30.0 | 53 |
川西云杉 P. likiangensis var. rubescens | 四川南部Southern Sichuan、青海南部Southern Qinghai、西藏东部Eastern Tibet | 94.4-103.0 | 28.3-32.9 | 33 |
林芝云杉 P. likiangensis var. linzhiensis | 西藏东南部Southeast Tibet、云南西北部Northwest Yunnan、四川西南部Southwest Sichuan | 92.9-100.6 | 27.8-30.3 | 29 |
紫果云杉P. purpurea | 四川北部Northern Sichuan、甘肃南部Southern Gansu、青海Qinghai | 101.1-104.1 | 30.1-35.8 | 47 |
须弥云杉P. spinulosa | 西藏南部Southern Tibet | 88.5-92.0 | 27.2-30.3 | 17 |
长叶云杉P. smithiana | 西藏南部Southern Tibet | 81.8-86.0 | 28.0-30.0 | 27 |
表2
云杉属主要树种地理分布区的气候因子描述性统计结果"
气候因子Climatic factors | 平均值 Mean | SD | 95%置信区间 95% confidence interval | 最小值 Minimum | 最大值 Maximum | CV (%) | |
下限 Lowerbound | 上限 Upperbound | ||||||
年均气温Annual mean temperature/℃ | 5.92 | 2.92 | 5.63 | 6.21 | -1.40 | 13.30 | 49.35 |
月均昼夜气温差Monthly mean temperature difference between day and night/℃ | 13.00 | 1.45 | 12.86 | 13.15 | 8.30 | 16.40 | 11.15 |
最暖月最高气温Maximal temperature of the warmest month/℃ | 21.38 | 3.72 | 21.01 | 21.76 | 15.00 | 32.10 | 17.39 |
最冷月最低气温Minimal temperature of the coldest month/℃ | -13.36 | 6.07 | -13.96 | -12.75 | -32.20 | -0.20 | 45.43 |
年均气温变幅Temperature annual range/℃ | 34.74 | 7.93 | 33.95 | 35.53 | 22.30 | 56.70 | 22.82 |
年均降水量Annual precipitation/mm | 691.70 | 187.10 | 673.10 | 710.40 | 277.00 | 1 310.00 | 27.10 |
最暖季降水量Precipitation of the warmest quarter/mm | 403.40 | 123.70 | 391.10 | 415.70 | 184.00 | 883.00 | 30.70 |
最冷季降水量Precipitation of the coldest quarter/mm | 16.20 | 7.40 | 15.50 | 16.90 | 4.00 | 41.00 | 45.90 |
干旱指数Aridity index | 17.74 | 4.36 | 17.30 | 18.17 | 7.94 | 29.63 | 24.59 |
表3
云杉属各树种地理分布区的气候因子特点①"
气候因子 Climatic factors | 项目 Item | 鱼鳞云杉 Picea jezoensis | 红皮云杉 P.koraiensis | 白杄 P.meyeri | 青杄 P.wilsonii | 云杉 P.asperata | 青海云杉 P.crassifolia | 丽江云杉 P.likiangensis var.likiangensis | 川西云杉 P.likiangensis var.rubescens | 林芝云杉 P.likiangensis var.linzhiensis | 紫果云杉 P.purpurea | 长叶云杉 P.smithiana | 须弥云杉 P.spinulosa |
年均气温 Annual mean temperature/℃ | 平均值Mean | 2.47a | 2.83ab | 5.20d | 7.99e | 5.24d | 3.66bc | 6.99d | 4.73cd | 7.70e | 3.97c | 9.86f | 9.76f |
SD | 1.70 | 0.89 | 2.93 | 3.15 | 1.73 | 1.71 | 1.80 | 1.20 | 1.33 | 1.77 | 1.51 | 0.98 | |
CV | 68.83 | 31.45 | 56.35 | 39.42 | 33.02 | 46.72 | 25.75 | 25.37 | 17.27 | 44.58 | 15.31 | 10.04 | |
月均昼夜气温差Monthly mean temperature difference between day and night/℃ | 平均值Mean | 12.52bc | 12.60bc | 12.56bc | 12.03b | 14.41d | 12.96c | 12.61bc | 14.30d | 12.70c | 14.38d | 11.14a | 12.61bc |
SD | 0.64 | 0.43 | 0.73 | 1.44 | 1.42 | 1.18 | 1.10 | 1.24 | 1.07 | 1.17 | 0.21 | 0.43 | |
CV | 5.11 | 3.41 | 5.81 | 11.97 | 9.85 | 9.10 | 8.72 | 8.67 | 8.43 | 8.14 | 1.89 | 3.41 | |
最暖月最高气温 Maximal temperature of warmest month/℃ | 平均值Mean | 25.62f | 25.81f | 25.83f | 26.14f | 20.40de | 20.17cde | 18.16a | 18.69ab | 19.94cde | 18.98abc | 19.68bcd | 21.00e |
SD | 1.54 | 1.29 | 2.96 | 3.68 | 1.56 | 1.81 | 1.58 | 1.57 | 1.64 | 1.74 | 1.57 | 1.06 | |
CV | 6.01 | 5.00 | 11.46 | 14.08 | 7.65 | 8.97 | 8.70 | 8.40 | 8.22 | 9.17 | 7.98 | 5.05 | |
最冷月最低气温 Minimal temperature of coldest month/℃ | 平均值Mean | -25.25a | -24.71a | -18.28b | -12.63e | -13.95de | -16.26c | -8.72f | -13.22e | -8.05f | -15.37cd | -3.19h | -6.36g |
SD | 2.81 | 1.67 | 3.49 | 4.41 | 2.67 | 2.16 | 2.64 | 1.74 | 1.78 | 2.74 | 1.75 | 1.04 | |
CV | 11.13 | 6.76 | 19.09 | 34.92 | 19.14 | 13.28 | 30.28 | 13.16 | 22.11 | 17.83 | 54.86 | 16.35 | |
年均气温变幅 Temperature annual range/℃ | 平均值Mean | 50.87h | 50.52h | 44.10g | 38.77f | 34.35d | 36.44e | 26.88b | 31.91c | 28.00b | 34.34d | 22.87a | 27.35b |
SD | 3.07 | 2.34 | 2.94 | 4.44 | 1.99 | 1.62 | 2.15 | 2.39 | 1.90 | 2.27 | 0.33 | 0.87 | |
CV | 6.03 | 4.63 | 6.67 | 11.45 | 5.79 | 4.45 | 8.00 | 7.49 | 6.79 | 6.61 | 1.44 | 3.18 | |
年均降水量 Annual precipitation/mm | 平均值Mean | 671.6cd | 656.30c | 469.40a | 536.40b | 700.60cd | 500.60ab | 837.50e | 655.90c | 692.40cd | 727.80d | 957.20f | 1 028.90g |
SD | 87.39 | 82.78 | 91.12 | 121.48 | 80.48 | 84.71 | 118.60 | 119.66 | 80.80 | 124.14 | 189.96 | 125.74 | |
CV | 13.01 | 12.61 | 19.41 | 22.65 | 11.49 | 16.92 | 14.16 | 18.24 | 11.67 | 17.06 | 19.85 | 12.22 | |
最暖季降水量 Precipitation of warmest quarter/mm | 平均值Mean | 414.10e | 407.60e | 312.40b | 305.30b | 348.80c | 268.70a | 518.30f | 385.30de | 412.20e | 370.80cd | 638.90g | 622.90g |
SD | 58.35 | 50.84 | 61.91 | 66.40 | 41.91 | 31.08 | 80.24 | 56.29 | 48.46 | 61.84 | 136.30 | 72.53 | |
CV | 14.09 | 12.47 | 19.82 | 21.75 | 12.02 | 11.57 | 15.48 | 14.61 | 11.76 | 16.68 | 21.33 | 11.64 | |
最冷季降水量 Precipitation of coldest quarter/mm | 平均值Mean | 24.10f | 23.70f | 12.00ab | 13.20bcd | 15.70cde | 9.30a | 16.50de | 12.80bc | 15.40cde | 16.40de | 31.10g | 18.30e |
SD | 6.09 | 5.93 | 4.01 | 4.76 | 3.49 | 2.57 | 7.33 | 5.35 | 8.01 | 4.95 | 2.90 | 8.82 | |
CV | 25.27 | 25.02 | 33.42 | 36.06 | 22.23 | 27.63 | 44.42 | 41.80 | 52.01 | 30.18 | 9.32 | 48.20 | |
干旱指数 Aridity index | 平均值Mean | 18.98cd | 18.34bcd | 12.37a | 13.08a | 18.31bcd | 13.66a | 20.93ef | 17.40bc | 17.01b | 19.71de | 22.29f | 24.05g |
SD | 2.61 | 2.43 | 2.78 | 2.72 | 1.82 | 2.24 | 2.69 | 3.25 | 1.85 | 3.43 | 4.12 | 2.80 | |
CV | 13.75 | 13.25 | 22.47 | 20.8 | 9.94 | 16.40 | 12.85 | 18.68 | 10.88 | 17.40 | 18.48 | 11.64 |
表4
云杉属各树种间9个气候因子的方差分析①"
气候因子 Climatic factors | 平方和 Sums of squares | 均方 Mean squares | F | 方差来源 Percentage of variance (%) | ||||
种间 Between species | 种内 Within species | 种间a Between species | 种内b Within species | 种间 Between species | 种内 Within species | |||
年均气温Annual mean temperature/℃ | 1 864.28 | 1 450.30 | 169.48 | 3.85 | 44.02*** | 56.24 | 43.76 | |
月均昼夜气温差Monthly mean temperature difference between day and night/℃ | 352.96 | 457.60 | 32.09 | 1.21 | 26.52*** | 43.55 | 56.45 | |
最暖月最高气温Maximal temperature of the warmest month/℃ | 3 695.99 | 1 667.16 | 336.00 | 4.42 | 76.02*** | 68.91 | 31.09 | |
最冷月最低气温Minimal temperature of the coldest month/℃ | 11 452.33 | 2 862.00 | 1 041.12 | 7.59 | 137.17*** | 80.01 | 19.99 | |
年均气温变幅Temperature annual range/℃ | 21 876.06 | 2 517.31 | 1 988.73 | 6.68 | 297.71*** | 89.68 | 10.32 | |
年均降水量Annual precipitation/mm | 8 831 772.37 | 4 748 824.41 | 802 888.40 | 12 596.35 | 63.74*** | 65.03 | 34.97 | |
最暖季降水量Precipitation of eht warmest quarter/mm | 4 298 306.29 | 1 635 479.34 | 390 755.12 | 4 338.14 | 90.07*** | 72.44 | 27.56 | |
最冷季降水量Precipitation of eht coldest quarter/mm | 9 326.79 | 12 114.97 | 847.89 | 32.14 | 26.38*** | 43.50 | 56.50 | |
干旱指数Aridity index | 4 482.29 | 2 901.45 | 407.48 | 7.70 | 52.92*** | 60.70 | 39.30 |
表5
各气候因子的重要性和显著性检验结果"
气候因子 Climatic factors | 重要性排序 Significance order | 解释量 Explained variation (%) | F | P |
年均气温Annual mean temperature/℃ | 5 | 20.8 | 102.87 | < 0.001 |
月均昼夜气温差Monthly mean temperature difference between day and night/℃ | 9 | 0.2 | 0.30 | 0.60 |
最暖月最高气温Maximal temperature of the warmest month/℃ | 3 | 40.2 | 262.44 | < 0.001 |
最冷月最低气温Minimal temperature of the coldest month/℃ | 2 | 66.8 | 785.47 | < 0.001 |
年均气温变幅Temperature annual range/℃ | 1 | 84.2 | 2 085.07 | < 0.001 |
年均降水量Annual precipitation/mm | 4 | 21.4 | 106.34 | < 0.001 |
最暖季降水量Precipitation of the warmest quarter/mm | 6 | 15.9 | 74.06 | < 0.001 |
最冷季降水量Precipitation of eht coldest quarter/mm | 8 | 0.1 | 0.94 | 0.33 |
干旱指数Aridity index | 7 | 13.1 | 58.98 | < 0.001 |
表6
中国云杉属树种地理分布区气候因子的RDA分析结果"
项目Item | 排序轴Ordination axis | |
轴1 Axis 1 | 轴2 Axis 2 | |
年均气温Annual mean temperature/℃ | -0.46 | -0.06 |
最暖月最高气温Maximal temperature of the warmest month/℃ | 0.64 | 0.06 |
最冷月最低气温Minimal temperature of the coldest month/℃ | -0.83 | -0.04 |
年均气温变幅Temperature annual range/℃ | 0.93 | 0.06 |
年均降水量Annual precipitation/mm | -0.46 | -0.38 |
最暖季降水量Precipitation of the warmest quarter/mm | -0.40 | -0.36 |
干旱指数Aridity index | -0.36 | -0.37 |
特征值Eigenvalues | 0.88 | 0.01 |
树种-环境相关性Species-environment correlations | 0.96 | 0.53 |
树种数据累积解释率Cumulative percentage variance of species data(%) | 88.2 | 89.2 |
树种-环境关系累积解释率Cumulative percentage variance of species-environment relation(%) | 99 | 100 |
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