秦岭中段不同坡向锐齿栎种子雨、土壤种子库与幼苗更新
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摘要
以秦岭中段不同坡向(阳坡、半阳坡、半阴坡及阴坡)锐齿栎天然林为对象,对锐齿栎种子雨、种子库的数量、质量动态以及幼苗发育过程进行研究.结果表明:锐齿栎种子雨一般从8月中旬开始,9月中下旬到10月上旬达到高峰期,11月中上旬结束;不同坡向的种子雨降落历程、发生时间和组成不同;半阳坡种子雨强度最大,为134.13粒·m~(-2),其余大小依次为阳坡、半阴坡、阴坡;阳坡种子雨降落时间最早、持续时间最长,高峰期持续时间也最长,阴坡种子雨降落时间最晚、持续时间最短,高峰期持续时间也最短;种子活力及成熟种子占其种子雨的比例大小顺序为半阳坡>阳坡>半阴坡>阴坡.从种子雨降落结束到翌年8月,各坡向土壤种子库总储量大小顺序均为半阳坡>阳坡>半阴坡>阴坡;各坡向的土壤种子库中成熟、未成熟、被啃食种子数量和种子活力均随时间变化呈递减趋势,而霉烂种子数量增加;各坡向土壤种子库中的种子均主要集中在枯落物层,其次为0～2 cm土层,而在2～5 cm土层中只有极少量种子存在.4种坡向中锐齿栎幼苗的密度差异显著,半阳坡幼苗最多,其余依次是阳坡、半阴坡和阴坡.半阳坡更适合锐齿栎种子的萌发和幼苗生长.
We studied the seed rain, seed pool quantity and quality dynamics, and seedling develop-ment of natural Quercus aliena var. acureserrata forests in different slopes(sunny, semi-sunny, semi-shady and shady slope) of the middle Qinling Mountains, China. The results showed that seed rain of Q. aliena var. acureserrata generally began at mid-August, peaked from mid-late September to early October, and ended in mid-early November. The dissemination process, occurrence time, and composition of seed rain in four slopes were different. The seed rain intensity of the semi-sunny slope was the highest, which was 134.13 seeds·m~(-2), followed by sunny slope, semi-shady slope, and shady slope. The seed rain on the sunny slope occurred first, and the duration of the whole and peak period were the longest. The seed rain on the shady slope occurred at the latest, and the duration of the whole and peak period were the shortest. The seed viability and mature seeds proportion were semi-sunny slope > sunny slope > semi-shady slope > shady slope. From the end of the seed rain to the August of the next year, the size of soil seed bank was semi-sunny slope > sunny slope > semi-shady slope > shady slope. The number of mature, immature and nibbled seeds and seed vigor of soil seed bank showed a decreasing trend with time, while the number of moldy seeds increased. The seeds in soil seed bank mainly existed in the litter layer, followed by the 0-2 cm soil layer, with few seeds in the 2-5 cm soil layer. The seedling density of Q. aliena var. acureserrata was significantly different among the four slopes. The semi-sunny slope, with the highest seedling density, was more suitable for seed germination and seedling growth of Q. aliena var. acureserrata than other slopes.
We studied the seed rain, seed pool quantity and quality dynamics, and seedling develop-ment of natural Quercus aliena var. acureserrata forests in different slopes(sunny, semi-sunny, semi-shady and shady slope) of the middle Qinling Mountains, China. The results showed that seed rain of Q. aliena var. acureserrata generally began at mid-August, peaked from mid-late September to early October, and ended in mid-early November. The dissemination process, occurrence time, and composition of seed rain in four slopes were different. The seed rain intensity of the semi-sunny slope was the highest, which was 134.13 seeds·m~(-2), followed by sunny slope, semi-shady slope, and shady slope. The seed rain on the sunny slope occurred first, and the duration of the whole and peak period were the longest. The seed rain on the shady slope occurred at the latest, and the duration of the whole and peak period were the shortest. The seed viability and mature seeds proportion were semi-sunny slope > sunny slope > semi-shady slope > shady slope. From the end of the seed rain to the August of the next year, the size of soil seed bank was semi-sunny slope > sunny slope > semi-shady slope > shady slope. The number of mature, immature and nibbled seeds and seed vigor of soil seed bank showed a decreasing trend with time, while the number of moldy seeds increased. The seeds in soil seed bank mainly existed in the litter layer, followed by the 0-2 cm soil layer, with few seeds in the 2-5 cm soil layer. The seedling density of Q. aliena var. acureserrata was significantly different among the four slopes. The semi-sunny slope, with the highest seedling density, was more suitable for seed germination and seedling growth of Q. aliena var. acureserrata than other slopes.
引文
[1] Nathan R,Helene C.Spatial patterns of seed dispersal,their determinants and consequences for recruitment.Trends in Ecology and Evolution,2000,15:278-285
[2] Harper JL.Population Biology of Plants.London:Academic Press,1977:256-263
[3] Benigno GR,Mulualem T,Guillermo CM,et al.Soil seed bank assembly following secondary succession on abandoned agricultural fields in Nicaragua.Journal of Forestry Research,2009,20:349-354
[4] Brearley FQ,Proctor J,Suriantata,et al.Reproductive phonology over a 10-year period in a lowland evergreen rain forest of central Borneo.Journal of Ecology,2007,95:828-839
[5] Liu Z-G (刘足根),Zhu J-J (朱教君),Yuan X-L (袁小兰),et al.On seed rain and soil seed bank of Larix olgensis in montane regions of eastern Liaoning Pro-vince,China.Acta Ecologica Sinica (生态学报),2007,27(2):579-587 (in Chinese)
[6] Eriksson O.Evolution of seed dispersal and recruitment in clonal plants.Oikos,1992,63:439-448
[7] Yang M-S (杨茂生),Li G-C (李广潮),Chen D-S (陈东升),et al.The Forest Classification and Main Felling Regeneration in the Qinling Mountains.Xi’an:Shaanxi Science & Technology Press,1994:46-73 (in Chinese)
[8] Dang K-L (党坤良),Lei R-D (雷瑞德).Water-source conservation and regulating efficacy of different forests at Huoditang forest region in the Qinling Mountains.Journal of Soil and Water Conservation (土壤侵蚀与水土保持学报),1995,1(1):79-84 (in Chinese)
[9] Chang J-G (常建国),Liu S-R (刘世荣),Shi Z-M (史作民),et al.Response of soil respiration to soil temperature and moisture regimes in the natural forest of Quercus aliena var.acuteserrata.Scientia Silvae Sinicae (林业科学),2006,42(12):21-27 (in Chinese)
[10] Liu G-Q (刘广全),Ni W-J (倪文进),Liu H-F (刘慧芳),et al.Eco-environment and nutrient accumulation of sharptooth oak stands in southern slope of Mt.Qinling.Chinese Journal of Applied Ecology (应用生态学报),2002,13(5):513-518 (in Chinese)
[11] Li Z (李哲),Dong N-N (董宁宁),Hou L (侯琳),et al.Characteristics of soil and litter organic carbon and total nitrogen in different age groups of Quercus aliena var.acuteserrata forests in the Qinling Mountains.Journal of Central South University of Forestry & Technology (中南林业科技大学学报),2017,37(12):127-132 (in Chinese)
[12] Chen Z-C (陈志成),Liu C (刘畅),Liu X-J (刘晓静),et al.Effects of irradiation and tree size on the carbon-water balances of Quercus aliena var.acuteserrata.Scientia Silvae Sinicae (林业科学),2017,53(9):18-25 (in Chinese)
[13] Liu C (刘畅),Sun P-S (孙鹏森),Liu S-R (刘世荣),et al.Leaf photosynthetic pigment seasonal dyna-mic of Quercus aliena var.acuteserrata and its spectral reflectance response under throughfall elimination.Chinese Journal of Applied Ecology (应用生态学报),2017,28(4):1077-1086 (in Chinese)
[14] Wu H (吴昊),Wang D-X (王得祥),Huang Q-P (黄青平),et al.Influence of environmental factors on species diversity of pine-oak mixed forest in the middle part of south Qinling Mountains.Journal of Northwest A&F University (Natural Science) (西北农林科技大学学报:自然科学版),2012,40(9):41-50 (in Chinese)
[15] Liu G-Q (刘广全),Zhao S-D (赵士洞),Wang H (王浩),et al.Seasonal variation of growth and nutrient contents for photosynthetic organ of the sharptooth oak stands.Acta Ecologica Sinica (生态学报),2001,21(6):883-889 (in Chinese)
[16] Zhang W-H (张文辉),Lu Y-C (卢彦昌),Zhou J-Y (周建云),et al.Population structure and dynamics of Quercus variabilis in different habitats on northern slope of Bashan Mountain.Scientia Silvae Sinicae (林业科学),2008,44(7):12-15 (in Chinese)
[17] Yue H-J (岳红娟),Tong C (仝川),Zhu J-M (朱锦懋),et al.Seed rain and soil seed bank of endangered Taxus chinensis var.mairei in Fujian,China.Acta Ecologica Sinica (生态学报),2010,30(16):4389-4400 (in Chinese)
[18] Yin H-J (尹华军),Cheng X-Y (程新颖),Lai T (赖挺),et al.Seed rain,soil seed bank and seedling regeneration in a 65-year Picea asperata plantation in subalpine coniferous,western Sichuan,China.Chinese Journal of Plant Ecology (植物生态学报),2011,35(1):35-44 (in Chinese)
[19] Tanouchi H,Sato T,Takeshita K.Comparative studies on acorn and seedling dynamics of four Quercus species in evergreen broad-leaved forest.Journal of Plant Research,1994,107:153-159
[20] Sork VL.Evolutionary ecology of mast-seeding intempe-rate and tropical oaks (Quercus spp.).Vegetatio,1993,107/108:133-147
[21] Liu T (刘彤),Zhou Z-Q (周志强).Seed rain and soil seed bank of Quercus mongolica populations.Journal of Northeast Forestry University (东北林业大学学报),2007,35(5):22-23 (in Chinese)
[22] Wang W,Ma KP,Liu CR.Seed shadow of Quercus liaotungensis in a broad leaved forest in Dongling Mountain.Acta Botanica Sinica,2000,42:195-202
[23] Willson MF.Dispersal mode,seed shadows,and colonization patterns.Plant Ecology,1993,107:261-280
[24] Fu H-G (傅焕光),Yu G-M (于光明).The Cultivation and Utilization of Quercus variabilis.Beijing:China Forestry Press,1986 (in Chinese)
[25] Horton JS,Wright JT.The wood rat as an ecological factor in southern California watersheds.Ecology,1944,25:341-351
[26] Tian R-S (田瑞松).The influence of site condition on the growth of Larix principis-rupprechtii.Morden Horticulture (现代园艺),2018(19):76-77 (in Chinese)
[27] Watt AS.On the causes of failure of natural regeneration in British oak woods.Journal of Ecology,1919,7:173-203
[28] Liu GX,Mao PS,Wang YW,et al.Effects of adult neighbour and gap size on seedling emergence and early growth of Bromus inermis Leyss.Ecological Research,2008,23:197-205
[29] Koutecká E,Lep J.Effect of light and moisture conditions and seed age on germination of three closely related Myosotis species.Folia Geobotanica,2009,44:109-130
[30] Zhan X,Li L,Cheng W.Restoration of Stipa krylovii steppes in Inner Mongolia of China:Assessment of seed banks and vegetation composition.Journal of Arid Environments,2007,68:298-307
[31] Zhou Z-Q (周志琼),Bao W-K (包维楷),Wu F-Z (吴福忠),et al.Capability and limitation of regeneration of Rosa hugonis and Rosa soufieana in the dry valley of the upper Minjiang River.Acta Ecologica Sinica (生态学报),2009,29(4):1931-1939 (in Chinese)
[32] Garcia D,Obseo JR,Martinez I.Spatial concordance between seed rain and seeding establishment in bird-dispersed trees:Does scale matter.Journal of Ecology,2005,93:693-704
[33] Zhang X-B (张希彪),Wang R-J (王瑞娟),Shangguan Z-P (上官周平).Dynamics of seed rain and soil seed bank in Pinus tabuliformis Carr.forests in eroded hilly loess regions of the Loess Plateau in China.Acta Ecologica Sinica (生态学报),2009,29(4):1877-1884 (in Chinese)
[34] Crawley MJ,Long CR.Alternate bearing,predator satia-tion and seedling recruitment in Quercus robur L.Journal of Ecology,1995,83:683-696
[35] Du XJ,Guo QF,Gao XM,et al.Seed rain,soil seed bank,seed loss and regeneration of Castanopsis fargesii (Fagaceae) in a subtropical evergreen broad-leaved forest.Forest Ecology and Management,2007,238:212-219
[36] Zhang X-L (张秀亮),Xu J-W (徐建伟),Shen H-L (沈海龙),et al.Animal predation and dispersal of Sorbus pohuashanensis fruits and seeds.Chinese Journal of Applied Ecology (应用生态学报),2010,21(10):2677-2683 (in Chinese)
[37] Chai L-J (柴良建),Liu F-G (刘富贵),Xiao W-M (肖琬蒙),et al.Spatial distribution characteristics of vegetation coverage in Renshou County.Sichuan Forestry Exploration and Design (四川林勘设计),2017(2):50-56 (in Chinese)
[38] Wang Z-L (王中磊),Gao X-M (高贤明).The predating behavior of rodents and its effects to acorns of Quercus aliena var.acuteserrata.Acta Ecologica Sinica (生态学报),2006,26(11):3533-3541 (in Chinese)
[39] Harper JL.Population Biology of Plants.London:Academic Press,1977:83-147
[40] Wu M (吴敏),Zhang W-H (张文辉),Zhou J-Y (周建云),et al.Dynamics of Quercus variabilis seed rain and soil seed bank in different habitats on the north slope of Qinling Mountains.Chinese Journal of Applied Ecology (应用生态学报),2011,22(11):2807-2814 (in Chinese)
[2] Harper JL.Population Biology of Plants.London:Academic Press,1977:256-263
[3] Benigno GR,Mulualem T,Guillermo CM,et al.Soil seed bank assembly following secondary succession on abandoned agricultural fields in Nicaragua.Journal of Forestry Research,2009,20:349-354
[4] Brearley FQ,Proctor J,Suriantata,et al.Reproductive phonology over a 10-year period in a lowland evergreen rain forest of central Borneo.Journal of Ecology,2007,95:828-839
[5] Liu Z-G (刘足根),Zhu J-J (朱教君),Yuan X-L (袁小兰),et al.On seed rain and soil seed bank of Larix olgensis in montane regions of eastern Liaoning Pro-vince,China.Acta Ecologica Sinica (生态学报),2007,27(2):579-587 (in Chinese)
[6] Eriksson O.Evolution of seed dispersal and recruitment in clonal plants.Oikos,1992,63:439-448
[7] Yang M-S (杨茂生),Li G-C (李广潮),Chen D-S (陈东升),et al.The Forest Classification and Main Felling Regeneration in the Qinling Mountains.Xi’an:Shaanxi Science & Technology Press,1994:46-73 (in Chinese)
[8] Dang K-L (党坤良),Lei R-D (雷瑞德).Water-source conservation and regulating efficacy of different forests at Huoditang forest region in the Qinling Mountains.Journal of Soil and Water Conservation (土壤侵蚀与水土保持学报),1995,1(1):79-84 (in Chinese)
[9] Chang J-G (常建国),Liu S-R (刘世荣),Shi Z-M (史作民),et al.Response of soil respiration to soil temperature and moisture regimes in the natural forest of Quercus aliena var.acuteserrata.Scientia Silvae Sinicae (林业科学),2006,42(12):21-27 (in Chinese)
[10] Liu G-Q (刘广全),Ni W-J (倪文进),Liu H-F (刘慧芳),et al.Eco-environment and nutrient accumulation of sharptooth oak stands in southern slope of Mt.Qinling.Chinese Journal of Applied Ecology (应用生态学报),2002,13(5):513-518 (in Chinese)
[11] Li Z (李哲),Dong N-N (董宁宁),Hou L (侯琳),et al.Characteristics of soil and litter organic carbon and total nitrogen in different age groups of Quercus aliena var.acuteserrata forests in the Qinling Mountains.Journal of Central South University of Forestry & Technology (中南林业科技大学学报),2017,37(12):127-132 (in Chinese)
[12] Chen Z-C (陈志成),Liu C (刘畅),Liu X-J (刘晓静),et al.Effects of irradiation and tree size on the carbon-water balances of Quercus aliena var.acuteserrata.Scientia Silvae Sinicae (林业科学),2017,53(9):18-25 (in Chinese)
[13] Liu C (刘畅),Sun P-S (孙鹏森),Liu S-R (刘世荣),et al.Leaf photosynthetic pigment seasonal dyna-mic of Quercus aliena var.acuteserrata and its spectral reflectance response under throughfall elimination.Chinese Journal of Applied Ecology (应用生态学报),2017,28(4):1077-1086 (in Chinese)
[14] Wu H (吴昊),Wang D-X (王得祥),Huang Q-P (黄青平),et al.Influence of environmental factors on species diversity of pine-oak mixed forest in the middle part of south Qinling Mountains.Journal of Northwest A&F University (Natural Science) (西北农林科技大学学报:自然科学版),2012,40(9):41-50 (in Chinese)
[15] Liu G-Q (刘广全),Zhao S-D (赵士洞),Wang H (王浩),et al.Seasonal variation of growth and nutrient contents for photosynthetic organ of the sharptooth oak stands.Acta Ecologica Sinica (生态学报),2001,21(6):883-889 (in Chinese)
[16] Zhang W-H (张文辉),Lu Y-C (卢彦昌),Zhou J-Y (周建云),et al.Population structure and dynamics of Quercus variabilis in different habitats on northern slope of Bashan Mountain.Scientia Silvae Sinicae (林业科学),2008,44(7):12-15 (in Chinese)
[17] Yue H-J (岳红娟),Tong C (仝川),Zhu J-M (朱锦懋),et al.Seed rain and soil seed bank of endangered Taxus chinensis var.mairei in Fujian,China.Acta Ecologica Sinica (生态学报),2010,30(16):4389-4400 (in Chinese)
[18] Yin H-J (尹华军),Cheng X-Y (程新颖),Lai T (赖挺),et al.Seed rain,soil seed bank and seedling regeneration in a 65-year Picea asperata plantation in subalpine coniferous,western Sichuan,China.Chinese Journal of Plant Ecology (植物生态学报),2011,35(1):35-44 (in Chinese)
[19] Tanouchi H,Sato T,Takeshita K.Comparative studies on acorn and seedling dynamics of four Quercus species in evergreen broad-leaved forest.Journal of Plant Research,1994,107:153-159
[20] Sork VL.Evolutionary ecology of mast-seeding intempe-rate and tropical oaks (Quercus spp.).Vegetatio,1993,107/108:133-147
[21] Liu T (刘彤),Zhou Z-Q (周志强).Seed rain and soil seed bank of Quercus mongolica populations.Journal of Northeast Forestry University (东北林业大学学报),2007,35(5):22-23 (in Chinese)
[22] Wang W,Ma KP,Liu CR.Seed shadow of Quercus liaotungensis in a broad leaved forest in Dongling Mountain.Acta Botanica Sinica,2000,42:195-202
[23] Willson MF.Dispersal mode,seed shadows,and colonization patterns.Plant Ecology,1993,107:261-280
[24] Fu H-G (傅焕光),Yu G-M (于光明).The Cultivation and Utilization of Quercus variabilis.Beijing:China Forestry Press,1986 (in Chinese)
[25] Horton JS,Wright JT.The wood rat as an ecological factor in southern California watersheds.Ecology,1944,25:341-351
[26] Tian R-S (田瑞松).The influence of site condition on the growth of Larix principis-rupprechtii.Morden Horticulture (现代园艺),2018(19):76-77 (in Chinese)
[27] Watt AS.On the causes of failure of natural regeneration in British oak woods.Journal of Ecology,1919,7:173-203
[28] Liu GX,Mao PS,Wang YW,et al.Effects of adult neighbour and gap size on seedling emergence and early growth of Bromus inermis Leyss.Ecological Research,2008,23:197-205
[29] Koutecká E,Lep J.Effect of light and moisture conditions and seed age on germination of three closely related Myosotis species.Folia Geobotanica,2009,44:109-130
[30] Zhan X,Li L,Cheng W.Restoration of Stipa krylovii steppes in Inner Mongolia of China:Assessment of seed banks and vegetation composition.Journal of Arid Environments,2007,68:298-307
[31] Zhou Z-Q (周志琼),Bao W-K (包维楷),Wu F-Z (吴福忠),et al.Capability and limitation of regeneration of Rosa hugonis and Rosa soufieana in the dry valley of the upper Minjiang River.Acta Ecologica Sinica (生态学报),2009,29(4):1931-1939 (in Chinese)
[32] Garcia D,Obseo JR,Martinez I.Spatial concordance between seed rain and seeding establishment in bird-dispersed trees:Does scale matter.Journal of Ecology,2005,93:693-704
[33] Zhang X-B (张希彪),Wang R-J (王瑞娟),Shangguan Z-P (上官周平).Dynamics of seed rain and soil seed bank in Pinus tabuliformis Carr.forests in eroded hilly loess regions of the Loess Plateau in China.Acta Ecologica Sinica (生态学报),2009,29(4):1877-1884 (in Chinese)
[34] Crawley MJ,Long CR.Alternate bearing,predator satia-tion and seedling recruitment in Quercus robur L.Journal of Ecology,1995,83:683-696
[35] Du XJ,Guo QF,Gao XM,et al.Seed rain,soil seed bank,seed loss and regeneration of Castanopsis fargesii (Fagaceae) in a subtropical evergreen broad-leaved forest.Forest Ecology and Management,2007,238:212-219
[36] Zhang X-L (张秀亮),Xu J-W (徐建伟),Shen H-L (沈海龙),et al.Animal predation and dispersal of Sorbus pohuashanensis fruits and seeds.Chinese Journal of Applied Ecology (应用生态学报),2010,21(10):2677-2683 (in Chinese)
[37] Chai L-J (柴良建),Liu F-G (刘富贵),Xiao W-M (肖琬蒙),et al.Spatial distribution characteristics of vegetation coverage in Renshou County.Sichuan Forestry Exploration and Design (四川林勘设计),2017(2):50-56 (in Chinese)
[38] Wang Z-L (王中磊),Gao X-M (高贤明).The predating behavior of rodents and its effects to acorns of Quercus aliena var.acuteserrata.Acta Ecologica Sinica (生态学报),2006,26(11):3533-3541 (in Chinese)
[39] Harper JL.Population Biology of Plants.London:Academic Press,1977:83-147
[40] Wu M (吴敏),Zhang W-H (张文辉),Zhou J-Y (周建云),et al.Dynamics of Quercus variabilis seed rain and soil seed bank in different habitats on the north slope of Qinling Mountains.Chinese Journal of Applied Ecology (应用生态学报),2011,22(11):2807-2814 (in Chinese)