2006—2017年长白山阔叶红松林木本植物种子雨的时空动态
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摘要
种子是森林更新的基础.为研究长白山阔叶红松林种子雨组成及时空变化,在长白山阔叶红松林25 hm~2样地内设置150个面积为0.5 m~2的收集器.自2006年5月至2017年12月收集252次,共收集到成熟、未成熟种子764299粒,隶属12科17属27种,其中,主林层树种13种704231粒,占总数的90%以上.种子最多的4个树种依次为紫椴、水曲柳、色木槭、假色槭,每年各个收集器都收集到这4个树种的种子;种子雨大爆发现象在各林层树种均存在,次林层和林下层较主林层树种滞后1～2年;各林层树种集中在秋季产生种子,各树种的种子雨都存在较大的时空变异性,相对而言,多数树种的空间变异性大于时间变异性;与科罗拉多岛(BCI)热带森林样地和浙江古田山亚热带常绿林样地种子产量的年际变异系数相比,长白山阔叶红松林种子产量年际变异系数较大,支持高纬度地区种子产量变异性高于低纬度地区的假说.
Seeds are the basis for forest regeneration. To examine the composition and spatio-temporal dynamics of seed rains, a total of 150 seed traps of 0.5 m~2 were installed in a 25 hm~2 broad-leaved Korean pine(Pinus koraiensis) mixed forest plot in Changbai Mountains. With a total of 252 collections from May 2006 to September 2017, we collected 764299 mature and immature seeds which were belonged to 27 species, 17 genera, and 12 families. More than 90% of all collected seeds(704231 seeds) were from 13 canopy species. Seeds of four tree species, including Tilia amurensis, Fraxinus mandschurica, Acer mono, and Acer pseudo-sieboldianum could be collected every year from each trap. Mast-seeding was found in every canopy layer, but it happened one to two years earlier in the overstorey layer than midstorey and understorey layer. Almost all species produced seeds in autumn, with considerable spatiotemporal variation. Generally, the spatial variation of seeds was larger than temporal variation. Compared with annual variation coefficient of seeds in tropical forest of the Barro Colorado Island(BCI) and subtropical evergreen forest in the Gutianshan, annual variation coefficient of seeds in Changbai Mountains was higher, which supported the hypothesis that annual variation in seed rains would be lower in the tropics than that in higher latitudes.
Seeds are the basis for forest regeneration. To examine the composition and spatio-temporal dynamics of seed rains, a total of 150 seed traps of 0.5 m~2 were installed in a 25 hm~2 broad-leaved Korean pine(Pinus koraiensis) mixed forest plot in Changbai Mountains. With a total of 252 collections from May 2006 to September 2017, we collected 764299 mature and immature seeds which were belonged to 27 species, 17 genera, and 12 families. More than 90% of all collected seeds(704231 seeds) were from 13 canopy species. Seeds of four tree species, including Tilia amurensis, Fraxinus mandschurica, Acer mono, and Acer pseudo-sieboldianum could be collected every year from each trap. Mast-seeding was found in every canopy layer, but it happened one to two years earlier in the overstorey layer than midstorey and understorey layer. Almost all species produced seeds in autumn, with considerable spatiotemporal variation. Generally, the spatial variation of seeds was larger than temporal variation. Compared with annual variation coefficient of seeds in tropical forest of the Barro Colorado Island(BCI) and subtropical evergreen forest in the Gutianshan, annual variation coefficient of seeds in Changbai Mountains was higher, which supported the hypothesis that annual variation in seed rains would be lower in the tropics than that in higher latitudes.
引文
[1] Nathan R,Muller-Landau HC.Spatial patterns of seed dispersal,their determinants and consequences for recruitment.Trends in Ecology and Evolution,2000,15:278-285
[2] Du Y-J (杜彦君),Ma K-P (马克平).Temporal and spatial variation of seedfall in a broad-leaved evergreen forest in Gutianshan Nature Reserve of Zhejiang Pro-vince,China.Chinese Journal of Plant Ecology (植物生态学报),2012,36(8):717-728 (in Chinese)
[3] Du Y-J (杜彦君),Ma K-P (马克平).Advancements and prospects in forest seed rain studies.Biodiversity Science (生物多样性),2012,20(1):94-107 (in Chinese)
[4] Yu S-L (于顺利),Lang N-J (郎南军),Peng M-J (彭明俊),et al.Research advances in seed rain.Chinese Journal of Ecology (生态学杂志),2007,26(10):1646-1652 (in Chinese)
[5] Schupp EW,Milleron T,Russo SE,et al.Dissemination limitation and the origin and maintenance of species-rich tropical forests.Seed Dispersal & Frugivory Ecology Evolution & Conservation,2002,98:233-246
[6] Clark JS,Macklin E,Wood L.Stages and spatial scales of recruitment limitation in southern Appalachian forests.Ecological Monographs,1998,68:213-235
[7] Zhang J (张健),Li B-H (李步杭),Hao Z-Q (郝占庆),et al.Composition and seasonal dynamics of seed rain in broad-leaved Korean pine (Pinus koraiensis) mixed forest,Changbai Mountain.Acta Ecologica Sinica (生态学报),2008,28(6):2245-2254 (in Chinese)
[8] Rossi S,Morin H,Laprise D,et al.Testing masting mechanisms of boreal forest species at different stand densities.Oikos,2012,121:665-674
[9] Rossi S,Morin H,Gionest F,et al.Inter-and intra-annual patterns of seed rain in the black spruce stands of Quebec,Canada.iForest:Biogeosciences and Forestry,2016,10:e1-e7
[10] 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)
[11] Alvarez-Buylla ER,Martínez-Ramos M.Seed bank versus seed rain in the regeneration of a tropical pioneer tree.Oecologia,1990,84:314-325
[12] Montesinos D,Verdú M.Masting uncoupling:Mast seeding does not follow all mast flowering episodes in a dioecious juniper tree.Oikos,2012,121:1725-1736
[13] Detto M,Wright SJ,Calderón O,et al.Resource acquisition and reproductive strategies of tropical forest in response to the El Niňo-Southern Oscillation.Nature Communications,2018,9:913
[14] Desoto L,Tutor D,Torices R,et al.Pre-dispersal predation effect on seed packaging strategies and seed viability.Oecologia,2016,180:1-12
[15] Ayako K,Teruyoshi N,Wajiro S.Seed dormancy and germination characteristics in relation to the regeneration of Acer pycnanthum,a vulnerable tree species in Japan.Journal of Forest Research,2015,20:160-166
[16] Pearse IS,Koenig WD,Kelly D.Mechanisms of mast seeding:Resources,weather,cues,and selection.New Phytologist,2016,212:546-562
[17] Kelly D.The evolutionary ecology of mast seeding.Trends in Ecology and Evolution,1994,9:465-470
[18] Kelly D,Sork VL.Mast seeding in perennial plants:Why,how,where?Annual Review of Ecology & Systematics,2002,33:427-447
[19] Wright SJ,Muller-Landau HC,Calderón O,et al.Annual and spatial variation in seedfall and seedling recruitment in a neotropical forest.Ecology,2005,86:848-860
[20] Chen XS,Li QJ.Patterns of plant sexual systems in subtropical evergreen broad-leaved forests in Ailao Mountains,SW China.Journal of Plant Ecology,2008,1:179-185
[21] Jia X (贾翔),Jin H (金慧),Zhao Y (赵莹),et al.Characteristics of climate change in Changbai Mountain from 1958 to 2015.Journal of Beihua University (Natural Science) (北华大学学报:自然科学版),2017,18(6):727-731 (in Chinese)
[22] Wang Y-Y(王芸芸).Reproductive Traits and Its Relationship with Spatial Pattern of Woody Plants in Broad-leaved Korean Pine (Pinus koraiensis) Mixed Forests in Changbai Mountains.PhD Thesis.Shenyang:Shenyang Institute of Applied Ecology,Chinese Academy of Sciences,2017 (in Chinese)
[23] Zhang J (张健),Li B-H (李步杭),Bai X-J (白雪娇),et al.Composition and interannual dynamics of tree seedlings in broad-leaved Korean pine (Pinus koraiensis) mixed forest in Changbai Mountain.Biodiversity Science (生物多样性),2009,17(4):385-396 (in Chinese)
[24] Yuan Z-Q (原作强),Li B-H (李步杭),Bai X-J (白雪娇),et al.Composition and seasonal dynamics of litter falls in a broad-leaved Korean pine (Pinus koraiensis) mixed forest in Changbai Mountains,Northeast China.Chinese Journal of Applied Ecology (应用生态学报),2010,21(8):2171-2178 (in Chinese)
[25] Chen X-M (陈香茗).Spatiotemporal Dynamics of Seed Rain and Seedling in Broad-leaved Korean Pine Mixed Forests in Changbai Mountain.Master Thesis.Beijing:Beijing Forestry University,2011 (in Chinese)
[26] Li B-H (李步杭).Dispersal Dynamics and Reproductive Strategy of Seed Rain in a Broad-leaved Korean Pine (Pinus koraiensis) Mixed Forests in Changbai Mountains.PhD Thesis.Shenyang:Shenyang Institute of Applied Ecology,Chinese Academy of Sciences,2013 (in Chinese)
[27] Kuang X (匡旭).Regeneration of Tree Seedlings in a Broad-leaved Korean Pine (Pinus koraiensis) Mixed Forests in Changbai Mountains.PhD Thesis.Shenyang:Shenyang Institute of Applied Ecology,Chinese Academy of Sciences,2017 (in Chinese)
[28] Hao Z-Q (郝占庆),Li B-H (李步杭),Zhang J (张健),et al.Broadleaved Korean pine (Pinus koraiensis) mixed forest plot in Changbaishan of China:Community composition and structure.Chinese Journal of Plant Ecology (植物生态学报),2008,32(2):238-250 (in Chinese)
[29] Wang X-G (王绪高),Hao Z-Q (郝占庆),Ye J (叶吉),et al.Relationship between species abundance and spatial distribution pattern of broad-leaved Korean pine (Pinus koraiensis) mixed forest in Changbai Mountains of China.Chinese Journal of Ecology (生态学杂志),2008,27(2):145-150 (in Chinese)
[30] Volis S.Correlated patterns of variation in phenology and seed production in populations of two annual grasses along an aridity gradient.Evolutionary Ecology,2007,21:381-393
[31] Bogdziewicz M,Steele MA,Marino S,et al.Correlated seed failure as an environmental veto to synchronize reproduction of masting plants.New Phytologist,2017,219:1-11
[32] Ran N,Katul GG,Bohrer G,et al.Mechanistic models of seed dispersal by wind.Theoretical Ecology,2011,4:113-132
[33] Levin SA,Muller-Landau HC,Nathan R,et al.The ecology and evolution of seed dispersal:A theoretical perspective.Annual Review of Ecology,Evolution and Systematics,2003,34:575-604
[2] Du Y-J (杜彦君),Ma K-P (马克平).Temporal and spatial variation of seedfall in a broad-leaved evergreen forest in Gutianshan Nature Reserve of Zhejiang Pro-vince,China.Chinese Journal of Plant Ecology (植物生态学报),2012,36(8):717-728 (in Chinese)
[3] Du Y-J (杜彦君),Ma K-P (马克平).Advancements and prospects in forest seed rain studies.Biodiversity Science (生物多样性),2012,20(1):94-107 (in Chinese)
[4] Yu S-L (于顺利),Lang N-J (郎南军),Peng M-J (彭明俊),et al.Research advances in seed rain.Chinese Journal of Ecology (生态学杂志),2007,26(10):1646-1652 (in Chinese)
[5] Schupp EW,Milleron T,Russo SE,et al.Dissemination limitation and the origin and maintenance of species-rich tropical forests.Seed Dispersal & Frugivory Ecology Evolution & Conservation,2002,98:233-246
[6] Clark JS,Macklin E,Wood L.Stages and spatial scales of recruitment limitation in southern Appalachian forests.Ecological Monographs,1998,68:213-235
[7] Zhang J (张健),Li B-H (李步杭),Hao Z-Q (郝占庆),et al.Composition and seasonal dynamics of seed rain in broad-leaved Korean pine (Pinus koraiensis) mixed forest,Changbai Mountain.Acta Ecologica Sinica (生态学报),2008,28(6):2245-2254 (in Chinese)
[8] Rossi S,Morin H,Laprise D,et al.Testing masting mechanisms of boreal forest species at different stand densities.Oikos,2012,121:665-674
[9] Rossi S,Morin H,Gionest F,et al.Inter-and intra-annual patterns of seed rain in the black spruce stands of Quebec,Canada.iForest:Biogeosciences and Forestry,2016,10:e1-e7
[10] 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)
[11] Alvarez-Buylla ER,Martínez-Ramos M.Seed bank versus seed rain in the regeneration of a tropical pioneer tree.Oecologia,1990,84:314-325
[12] Montesinos D,Verdú M.Masting uncoupling:Mast seeding does not follow all mast flowering episodes in a dioecious juniper tree.Oikos,2012,121:1725-1736
[13] Detto M,Wright SJ,Calderón O,et al.Resource acquisition and reproductive strategies of tropical forest in response to the El Niňo-Southern Oscillation.Nature Communications,2018,9:913
[14] Desoto L,Tutor D,Torices R,et al.Pre-dispersal predation effect on seed packaging strategies and seed viability.Oecologia,2016,180:1-12
[15] Ayako K,Teruyoshi N,Wajiro S.Seed dormancy and germination characteristics in relation to the regeneration of Acer pycnanthum,a vulnerable tree species in Japan.Journal of Forest Research,2015,20:160-166
[16] Pearse IS,Koenig WD,Kelly D.Mechanisms of mast seeding:Resources,weather,cues,and selection.New Phytologist,2016,212:546-562
[17] Kelly D.The evolutionary ecology of mast seeding.Trends in Ecology and Evolution,1994,9:465-470
[18] Kelly D,Sork VL.Mast seeding in perennial plants:Why,how,where?Annual Review of Ecology & Systematics,2002,33:427-447
[19] Wright SJ,Muller-Landau HC,Calderón O,et al.Annual and spatial variation in seedfall and seedling recruitment in a neotropical forest.Ecology,2005,86:848-860
[20] Chen XS,Li QJ.Patterns of plant sexual systems in subtropical evergreen broad-leaved forests in Ailao Mountains,SW China.Journal of Plant Ecology,2008,1:179-185
[21] Jia X (贾翔),Jin H (金慧),Zhao Y (赵莹),et al.Characteristics of climate change in Changbai Mountain from 1958 to 2015.Journal of Beihua University (Natural Science) (北华大学学报:自然科学版),2017,18(6):727-731 (in Chinese)
[22] Wang Y-Y(王芸芸).Reproductive Traits and Its Relationship with Spatial Pattern of Woody Plants in Broad-leaved Korean Pine (Pinus koraiensis) Mixed Forests in Changbai Mountains.PhD Thesis.Shenyang:Shenyang Institute of Applied Ecology,Chinese Academy of Sciences,2017 (in Chinese)
[23] Zhang J (张健),Li B-H (李步杭),Bai X-J (白雪娇),et al.Composition and interannual dynamics of tree seedlings in broad-leaved Korean pine (Pinus koraiensis) mixed forest in Changbai Mountain.Biodiversity Science (生物多样性),2009,17(4):385-396 (in Chinese)
[24] Yuan Z-Q (原作强),Li B-H (李步杭),Bai X-J (白雪娇),et al.Composition and seasonal dynamics of litter falls in a broad-leaved Korean pine (Pinus koraiensis) mixed forest in Changbai Mountains,Northeast China.Chinese Journal of Applied Ecology (应用生态学报),2010,21(8):2171-2178 (in Chinese)
[25] Chen X-M (陈香茗).Spatiotemporal Dynamics of Seed Rain and Seedling in Broad-leaved Korean Pine Mixed Forests in Changbai Mountain.Master Thesis.Beijing:Beijing Forestry University,2011 (in Chinese)
[26] Li B-H (李步杭).Dispersal Dynamics and Reproductive Strategy of Seed Rain in a Broad-leaved Korean Pine (Pinus koraiensis) Mixed Forests in Changbai Mountains.PhD Thesis.Shenyang:Shenyang Institute of Applied Ecology,Chinese Academy of Sciences,2013 (in Chinese)
[27] Kuang X (匡旭).Regeneration of Tree Seedlings in a Broad-leaved Korean Pine (Pinus koraiensis) Mixed Forests in Changbai Mountains.PhD Thesis.Shenyang:Shenyang Institute of Applied Ecology,Chinese Academy of Sciences,2017 (in Chinese)
[28] Hao Z-Q (郝占庆),Li B-H (李步杭),Zhang J (张健),et al.Broadleaved Korean pine (Pinus koraiensis) mixed forest plot in Changbaishan of China:Community composition and structure.Chinese Journal of Plant Ecology (植物生态学报),2008,32(2):238-250 (in Chinese)
[29] Wang X-G (王绪高),Hao Z-Q (郝占庆),Ye J (叶吉),et al.Relationship between species abundance and spatial distribution pattern of broad-leaved Korean pine (Pinus koraiensis) mixed forest in Changbai Mountains of China.Chinese Journal of Ecology (生态学杂志),2008,27(2):145-150 (in Chinese)
[30] Volis S.Correlated patterns of variation in phenology and seed production in populations of two annual grasses along an aridity gradient.Evolutionary Ecology,2007,21:381-393
[31] Bogdziewicz M,Steele MA,Marino S,et al.Correlated seed failure as an environmental veto to synchronize reproduction of masting plants.New Phytologist,2017,219:1-11
[32] Ran N,Katul GG,Bohrer G,et al.Mechanistic models of seed dispersal by wind.Theoretical Ecology,2011,4:113-132
[33] Levin SA,Muller-Landau HC,Nathan R,et al.The ecology and evolution of seed dispersal:A theoretical perspective.Annual Review of Ecology,Evolution and Systematics,2003,34:575-604