青藏高原东北缘3种灌木数量性状、龄级结构及自疏特征
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摘要
以青藏高原东北缘高寒灌丛优势种金露梅(Potentilla fruticosa)、高山柳(Salix cupularis)和头花杜鹃(Rhododendron capitatum)为对象,通过对其数量性状(株高、冠幅直径、分枝数、单株重、个体分枝重)与年龄结构及自疏特征分析,构建3种灌木株丛生物量估测模型,明确其年龄结构模式,确定3种灌木分枝数和分枝重之间是否存在自疏特征。结果表明:1)植物种群数量性状对其地上部分单株重影响为金露梅>高山柳>头花杜鹃;2)冠幅直径可作为3种灌木单株重估测模型适宜指标,基于冠幅直径(CD,cm)构建的灌木单株重(PW, kg)估测模型PW_(金露梅)=–0.310+0.014CD,PW_(高山柳)=–1.688+0.033CD和PW_(头花杜鹃)=–0.203+0.013CD;3)头花杜鹃和高山柳属于增长型种群,具一定稳定性,金露梅属于稳定型种群;4)头花杜鹃株丛分枝数和分枝重之间存在明显自疏特征。研究认为,3种灌木的数量性状、年龄结构和自疏特征存在分异特征,是其对高寒生境的长期适应性结果。
The quantitative traits(plant height, crown diameter, number of branches, plant weight and branch weight), age structure and self-thinning rule of Potentilla fruticosa, Salix cupularis and Rhododendron capitatum on the northeast margin of the Qinghai-Tibet Plateau were studied. We constructed the estimation model of plant biomass of this three shrubs,determined their age structures, and clarified the self-thinning relationship between the number of branches and branch weight for each shrub. The results showed that: 1) The impact on the plant weight by the quantitative characteristics of each species was Potentilla fruticosa > Salix cupularis > Rhododendron capitatum; 2) The crown diameter could be used as a suitable index for estimating the shrub plant weight, the estimation models of plant weight(PW, kg) basing on the crown diameters(CD, cm) were PW = –0.310 + 0.014 CD for P. fruticosa, PW = –1.688 + 0.033 CD for S. cupularis and PW =–0.203 + 0.013 CD for R. capitatum, respectively; 3) S. cupularis and R. capitatum was a growing population and had a certain stability, while the P. fruticosa belonged to a stable population; 4) There was an obvious self-thinning law between branch weight and number of branches of R. capitatum. This research shows that the quantitative traits, age structure and self-thinning law for each species is different, which suggests that it is an adaptive result to different habitats and nutrient resources.
The quantitative traits(plant height, crown diameter, number of branches, plant weight and branch weight), age structure and self-thinning rule of Potentilla fruticosa, Salix cupularis and Rhododendron capitatum on the northeast margin of the Qinghai-Tibet Plateau were studied. We constructed the estimation model of plant biomass of this three shrubs,determined their age structures, and clarified the self-thinning relationship between the number of branches and branch weight for each shrub. The results showed that: 1) The impact on the plant weight by the quantitative characteristics of each species was Potentilla fruticosa > Salix cupularis > Rhododendron capitatum; 2) The crown diameter could be used as a suitable index for estimating the shrub plant weight, the estimation models of plant weight(PW, kg) basing on the crown diameters(CD, cm) were PW = –0.310 + 0.014 CD for P. fruticosa, PW = –1.688 + 0.033 CD for S. cupularis and PW =–0.203 + 0.013 CD for R. capitatum, respectively; 3) S. cupularis and R. capitatum was a growing population and had a certain stability, while the P. fruticosa belonged to a stable population; 4) There was an obvious self-thinning law between branch weight and number of branches of R. capitatum. This research shows that the quantitative traits, age structure and self-thinning law for each species is different, which suggests that it is an adaptive result to different habitats and nutrient resources.
引文
[1]张贞明,韩天虎.几种高寒灌丛地上植物量的估测模型.草业科学,2008, 25(1):10-13.ZHANG Z M, HAN T H. A mathematical model for aboveground biomass the in alpine shrub in Gannan, Gansu Province.Pratacultural Science, 2008, 25(1):10-13.
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[7]陶冶,张元明.荒漠灌木生物量多尺度估测:以梭梭为例.草业学报,2013, 22(6):1-10.TAO Y, ZHANG Y M. Multi-scale biomass estimation of desert shrubs:A case study of Haloxylon ammodendron in the Gurbantunggut Desert, China. Acta Prataculturae Sinica, 2013, 22(6):1-10.
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[9]杨宪龙,魏孝荣,邵明安.黄土高原北部典型灌丛枝条生物量估算模型.应用生态学报,2016, 27(10):3164-3172.YANG X L, WEI X R, SHAO M A. Stem biomass estimation models for dominant shrubs on the northern Loess Plateau of China.Journal of Applied Ecology, 2016, 27(10):3164-3172.
[10]邢福,郭继勋,魏春雁.退化草原狼毒个体年龄判定方法及其种群年龄结构的研究.应用生态学报,2004(11):2104-2108.XING F, GUO J X, WEI C Y. Judging method of individual age and age structure of Stellera chamaejasme population in degraded steppe. Journal of Applied Ecology, 2004(11):2104-2108.
[11]陈小勇,张庆费,吴化前,宋永昌.黄山西坡青冈种群结构与分布格局研究.生态学报,1996, 16(3):325-327.CHEN X Y, ZHANG Q F, WU H Q, SONG Y C. Study on the structure and spatial patterns of Cyclobalanopsis glauca populations in western Huangshan. Chinese Journal of Ecology, 1996, 16(3):325-327.
[12]李昌龙,马瑞君,王继和,李爱德,严子柱,李得禄.甘肃民勤连古城自然保护区优势种种群结构和动态研究.西北植物学报,2005(08):1628-1636.LI C L, MA R J, WANG J H, LI A D, YAN Z Z, LI D L. Community structures and dynamics of dominant species in Minqinliangucheng Nature Reserve of Gansu Province. Acta Botanica Boreali-Occidentalia Sinica, 2005(08):1628-1636.
[13]韩文轩,方精云.植物种群的自然稀疏规律-3/2还是-4/3.北京大学学报,2008,44(4):661-668.HAN W X, FANG J Y. Self-thinning law in plant populations:-3/2 vs-4/3. Journal of Beijing University, 2008, 44(4):661-668.
[14]周威,龙成,杨小波,龙文兴,李东海,卜广发.海南铜鼓岭灌木林稀疏规律.生态学报,2013, 33(20):6569-6576.ZHOU W, LONG C, YANG X B, LONG W X, LI D H, BO G F. The thinning regular of the shrubbery at Tongguling National Nature Reserve on Hainan Island, China. Chinese Journal of Ecology, 2013, 33(20):6569-6576.
[15]赵锦梅,赵品忠,耿妍,权佳,徐长林.祁连山东段不同高寒灌丛草地土壤性状特征变化.草地学报,2014, 22(5):991-997.ZHAO J M, ZHAO J Z, GENG Y, QUAN J, XU C L. Soil properties of different alpine shrub grasslands in eastern Qilian mountains. Acta Agrestia Sinica, 2014, 22(5):991-997.
[16]韦应莉,曹文侠,李建宏,张爱梅,李小龙.不同放牧与围封高寒灌丛草地土壤微生物群落结构PLFA分析.生态学报,2018,38(13):4897-4908.WEI Y L, CAO W X, LI J H,ZHANG A M,LI X L. Phospholipid fatty acid(PLFA)analysis of soil microbial community structure with different intensities of grazing and fencing in alpine shrubland. Chinese Journal of Ecology, 2018, 38(13):4897-4908.
[17]盛海彦,曹广民,李国荣,周靖靖,焦文月,李吉鹏,张平.放牧干扰对祁连山高寒金露梅灌丛草甸群落的影响.生态环境学报,2009, 18(1):235-241.SHENG H Y, CAO G M, LI G R, ZHOU J J, JIAO W Y, LI J P, ZHANG P. Effect of grazing disturbance on plant community of alpine meadow dominated by Potentilla fruticosa shrub on Qilian Mountain. Ecology and Environmental Sciences, 2009, 18(1):235-241.
[18]杨路存,赵玉红,徐文华,周国英.青海省高寒灌丛物种多样性、生物量及其关系.生态学报,2018, 38(1):309-315.YANG L C, ZHAO Y H, XU W H, ZHOU G Y. Species diversity, biomass, and their relationship in the alpine shrubberies of Qinghai Province. Chinese Journal of Ecology, 2018, 38(1):309-315.
[19]徐世晓,赵亮,赵新全,李英年,古松.青藏高原高寒灌丛非生长季节CO2通量特征.西北植物学报,2006(12):2528-2532.XU S X, ZHAO L, ZHAO X Q, LI Y N, GU S. Carbon dioxide flux characteristics of alpine shrubs in Qinghai-Tibet Plateau beyond the growing season. Acta Botanica Boreali-Occidentalia Sinica, 2006(12):2528-2532.
[20]王立红,王悦,王德凯,解李娜,马成仓.放牧对小叶锦鸡儿种群数量特征的影响.草业科学,2018, 35(4):800-806.WANG L H, WANG Y, WANG D K, XIE L N, MA C C. Effect of grazing on quantitative characteristics of Caragana microphylla population. Pratacultural Science, 2018, 35(4):800-806.
[21]徐长林.坡向对青藏高原东北缘高寒草甸植被构成和养分特征的影响.草业学报,2016, 25(4):26-35.XU C L. Variation in vegetation composition and nutrient characteristics related to aspect in an alpine meadow in the notheast margin of the Qinghai-Tibet Plateau. Acta Prataculturae Sinica, 2016, 25(4):26-35.
[22]龙瑞军,徐长林,胡自治,丁文广,张映生,康天福.天祝高山草原15种饲用灌木的热值及季节动态.生态学杂志,1993, 12(5):13-16.LONG R J, XU C L, HU Z Z, DING W G, ZHANG Y S, KANG T F. Calorific value and its seasonal dynamics of fodder shrub species on Tianzhu Alpine grasslands. Chinese Journal of Ecology, 1993, 12(5):13-16.
[23]张德罡,曹文侠,蒲小鹏,胡自治.东祁连山杜鹃属植物形态特征及其生态适应性.草原与草坪,2003(1):27-30, 37.ZHANG D G, CAO W X, PU X P, HU Z Z. Botanical morpha and ecological adaptation of rhododendron in eastern Qilian Mountains. Grassland and Turf, 2003(1):27-30, 37.
[24]于海英,许建初.气候变化对青藏高原植被影响研究综述.生态学杂志,2009, 28(4):747-754.YU H Y, XU J C. Effects of climate change on vegetation on Qinghai-Tibet Plateau. Chinese Journal of Ecology, 2009, 28(4):747-754.
[25]张笑鹤,张远东,顾峰雪,刘世荣.西南地区灌丛归一化植被指数动态及其与气候因子的相关性.生态学杂志,2011,30(11):2577-2583.ZHANG X H, ZHANG Y D, GU F X, LIU S R. Dynamics of shrubs normalized difference vegetation index and its correlations with climatic factors in Southwest China. Chinese Journal of Ecology, 2011, 30(11):2577-2583.
[26]袁宏波,张锦春,褚建民,刘淑娟,唐进年,丁峰.库姆塔格沙漠典型植物种群年龄结构特征.西北植物学报,2011,31(11):2304-2309.YUAN H B, ZHANG J C, CHU J M, LIU S J, TANG J N, DING F. Characteristics of the age structures of three typical plant populations in Kumtag desert. Acta Botanica Boreali-Occidentalia Sinica, 2011, 31(11):2304-2309.
[2]曹文侠,徐长林,张德罡,师尚礼,姚拓.杜鹃灌丛草地土壤容重与水分特征对不同休牧模式的响应.草业学报,2011,20(3):28-35.CAO W X, XU C L, ZHANG D G, SHI S L, YAO T. Ecological responses of soil bulk density and water content to different nongrazing patterns in alpine rhododendron shrubland. Acta Prataculturae Sinica, 2011, 20(3):28-35.
[3]杨路存,李长斌,宁祎,聂秀青,徐文华,周国英.青海高寒金露梅灌丛碳密度及其分配格局.植物生态学报,2017, 41(1):62-70.YANG L C, LI C B, NING W, NIE X Q, XU W H, ZHOU G Y. Carbon density and its spatial distribution in the Potentilla fruticosa dominated alpine shrub in Qinghai, China. Journal of Plant Ecology, 2017, 41(1):62-70.
[4] LOREAU M. Biodiversity and ecosystem functioning:recent theoretical advances. Oikos, 2000, 91(1):3-17.
[5]杨弦,郭焱培,安尼瓦尔·买买提,刘鸿雁,马文红,于顺利,唐志尧.中国北方温带灌从生物量的分布及其与环境的关系.植物生态学报,2017, 41(1):22-30.YANG X, GUO Y P, MOHHAMOT A, LIU H Y, MA W H, YU S L, TANG Z Y. Distribution of biomass in relation to environments in shrublands of temperate China. Journal of Plant Ecology, 2017, 41(1):22-30.
[6]曹文侠,李文.千里香杜鹃根系生物量时空动态特征及其生态适应性.草业学报,2016, 25(7):52-61.CAO W X, LI W. Spatio-temporal trends for fine root biomass of alpine Rhododendron thymifolium and their significance for ecological adaptation in Qilian Mountains. Acta Prataculturae Sinica, 2016, 25(7):52-61.
[7]陶冶,张元明.荒漠灌木生物量多尺度估测:以梭梭为例.草业学报,2013, 22(6):1-10.TAO Y, ZHANG Y M. Multi-scale biomass estimation of desert shrubs:A case study of Haloxylon ammodendron in the Gurbantunggut Desert, China. Acta Prataculturae Sinica, 2013, 22(6):1-10.
[8] SALIS S M, ASSIS M A, MATTOS P P, PIAO A C S. Estimating the aboveground biomass and wood volume of savanna woodlands in Brazil's Pantanal wetlands based on allometric correlations. Forest Ecology and Management, 2006, 228:61-68.
[9]杨宪龙,魏孝荣,邵明安.黄土高原北部典型灌丛枝条生物量估算模型.应用生态学报,2016, 27(10):3164-3172.YANG X L, WEI X R, SHAO M A. Stem biomass estimation models for dominant shrubs on the northern Loess Plateau of China.Journal of Applied Ecology, 2016, 27(10):3164-3172.
[10]邢福,郭继勋,魏春雁.退化草原狼毒个体年龄判定方法及其种群年龄结构的研究.应用生态学报,2004(11):2104-2108.XING F, GUO J X, WEI C Y. Judging method of individual age and age structure of Stellera chamaejasme population in degraded steppe. Journal of Applied Ecology, 2004(11):2104-2108.
[11]陈小勇,张庆费,吴化前,宋永昌.黄山西坡青冈种群结构与分布格局研究.生态学报,1996, 16(3):325-327.CHEN X Y, ZHANG Q F, WU H Q, SONG Y C. Study on the structure and spatial patterns of Cyclobalanopsis glauca populations in western Huangshan. Chinese Journal of Ecology, 1996, 16(3):325-327.
[12]李昌龙,马瑞君,王继和,李爱德,严子柱,李得禄.甘肃民勤连古城自然保护区优势种种群结构和动态研究.西北植物学报,2005(08):1628-1636.LI C L, MA R J, WANG J H, LI A D, YAN Z Z, LI D L. Community structures and dynamics of dominant species in Minqinliangucheng Nature Reserve of Gansu Province. Acta Botanica Boreali-Occidentalia Sinica, 2005(08):1628-1636.
[13]韩文轩,方精云.植物种群的自然稀疏规律-3/2还是-4/3.北京大学学报,2008,44(4):661-668.HAN W X, FANG J Y. Self-thinning law in plant populations:-3/2 vs-4/3. Journal of Beijing University, 2008, 44(4):661-668.
[14]周威,龙成,杨小波,龙文兴,李东海,卜广发.海南铜鼓岭灌木林稀疏规律.生态学报,2013, 33(20):6569-6576.ZHOU W, LONG C, YANG X B, LONG W X, LI D H, BO G F. The thinning regular of the shrubbery at Tongguling National Nature Reserve on Hainan Island, China. Chinese Journal of Ecology, 2013, 33(20):6569-6576.
[15]赵锦梅,赵品忠,耿妍,权佳,徐长林.祁连山东段不同高寒灌丛草地土壤性状特征变化.草地学报,2014, 22(5):991-997.ZHAO J M, ZHAO J Z, GENG Y, QUAN J, XU C L. Soil properties of different alpine shrub grasslands in eastern Qilian mountains. Acta Agrestia Sinica, 2014, 22(5):991-997.
[16]韦应莉,曹文侠,李建宏,张爱梅,李小龙.不同放牧与围封高寒灌丛草地土壤微生物群落结构PLFA分析.生态学报,2018,38(13):4897-4908.WEI Y L, CAO W X, LI J H,ZHANG A M,LI X L. Phospholipid fatty acid(PLFA)analysis of soil microbial community structure with different intensities of grazing and fencing in alpine shrubland. Chinese Journal of Ecology, 2018, 38(13):4897-4908.
[17]盛海彦,曹广民,李国荣,周靖靖,焦文月,李吉鹏,张平.放牧干扰对祁连山高寒金露梅灌丛草甸群落的影响.生态环境学报,2009, 18(1):235-241.SHENG H Y, CAO G M, LI G R, ZHOU J J, JIAO W Y, LI J P, ZHANG P. Effect of grazing disturbance on plant community of alpine meadow dominated by Potentilla fruticosa shrub on Qilian Mountain. Ecology and Environmental Sciences, 2009, 18(1):235-241.
[18]杨路存,赵玉红,徐文华,周国英.青海省高寒灌丛物种多样性、生物量及其关系.生态学报,2018, 38(1):309-315.YANG L C, ZHAO Y H, XU W H, ZHOU G Y. Species diversity, biomass, and their relationship in the alpine shrubberies of Qinghai Province. Chinese Journal of Ecology, 2018, 38(1):309-315.
[19]徐世晓,赵亮,赵新全,李英年,古松.青藏高原高寒灌丛非生长季节CO2通量特征.西北植物学报,2006(12):2528-2532.XU S X, ZHAO L, ZHAO X Q, LI Y N, GU S. Carbon dioxide flux characteristics of alpine shrubs in Qinghai-Tibet Plateau beyond the growing season. Acta Botanica Boreali-Occidentalia Sinica, 2006(12):2528-2532.
[20]王立红,王悦,王德凯,解李娜,马成仓.放牧对小叶锦鸡儿种群数量特征的影响.草业科学,2018, 35(4):800-806.WANG L H, WANG Y, WANG D K, XIE L N, MA C C. Effect of grazing on quantitative characteristics of Caragana microphylla population. Pratacultural Science, 2018, 35(4):800-806.
[21]徐长林.坡向对青藏高原东北缘高寒草甸植被构成和养分特征的影响.草业学报,2016, 25(4):26-35.XU C L. Variation in vegetation composition and nutrient characteristics related to aspect in an alpine meadow in the notheast margin of the Qinghai-Tibet Plateau. Acta Prataculturae Sinica, 2016, 25(4):26-35.
[22]龙瑞军,徐长林,胡自治,丁文广,张映生,康天福.天祝高山草原15种饲用灌木的热值及季节动态.生态学杂志,1993, 12(5):13-16.LONG R J, XU C L, HU Z Z, DING W G, ZHANG Y S, KANG T F. Calorific value and its seasonal dynamics of fodder shrub species on Tianzhu Alpine grasslands. Chinese Journal of Ecology, 1993, 12(5):13-16.
[23]张德罡,曹文侠,蒲小鹏,胡自治.东祁连山杜鹃属植物形态特征及其生态适应性.草原与草坪,2003(1):27-30, 37.ZHANG D G, CAO W X, PU X P, HU Z Z. Botanical morpha and ecological adaptation of rhododendron in eastern Qilian Mountains. Grassland and Turf, 2003(1):27-30, 37.
[24]于海英,许建初.气候变化对青藏高原植被影响研究综述.生态学杂志,2009, 28(4):747-754.YU H Y, XU J C. Effects of climate change on vegetation on Qinghai-Tibet Plateau. Chinese Journal of Ecology, 2009, 28(4):747-754.
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