啮齿动物对秦岭松栎混交林建群种种子扩散格局的影响
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摘要
森林群落中的啮齿动物对林木种子存在着一定的取食偏好性,这种偏好性会使啮齿动物形成不同的捕食和贮藏策略,从而导致林木种子形成不同的扩散格局。以秦岭中段松栎混交林建群种锐齿槲栎、油松、华山松为研究对象,采用塑料标签标记种子方法,研究啮齿动物对种子传播和扩散格局的影响。结果表明:(1)种子特征是影响啮齿动物对其进行扩散的重要因子,油松种子以其质量小、种皮薄等特点吸引啮齿动物大量捕食,其原地取食率达到83.33%,显著高于华山松和锐齿槲栎种子。(2)锐齿槲栎和华山松种子大部分被搬运一定距离后再被取食和埋藏,其中,扩散距离在1 m以内、1—3 m和5—8 m的锐齿槲栎种子分别占59.12%、18.23%和13.26%,最远扩散距离达12 m处;华山松种子扩散距离主要介于在1—3 m(37.85%)和3—5 m(23.73%),在距离5—8 m(13.56%)、8—10 m(11.86%)和大于10 m(11.30%)区间也有一定的分布,最大扩散距离为12 m。(3)啮齿动物倾向于将锐齿槲栎种子搬运至松林内取食,而将华山松种子搬运至栎林内埋藏,说明不同林分类型及其环境因素对林木种子扩散后的分布格局具有重要影响。(4)3种类型种子被啮齿动物捕食的比例以油松最大(96.90%),锐齿槲栎次之(73.57%),华山松最小(50%);次年调查时,未被捕食的种子大部分已经被取食,仅有极少数锐齿槲栎种子萌发成幼苗(1.67%)。啮齿动物的捕食和贮藏行为对林木种子扩散及其成功更新都至关重要。
Rodents in forest communities have a feeding preference for tree seeds. This preference will lead to different predation and storage strategies for rodents, resulting in different diffusion patterns of tree seeds. In this study, the seed fate of Quercus aliena var. acuteserrata, Pinus armandii, and Pinus tabuliformis was studied to analyze the seed dispersal patterns of constructive species in the pine-oak mixed forests of the Qinling Mountains, Shaanxi Province, China. The results demonstrated that seed traits are the main factor affecting their removal by rodents. The in-situ seed predation rate of P. tabuliformis seeds was 83.33%, which was notably higher than that of other seed types, indicating that rodents preferentially fed on the small P. tabuliformis seeds. Q. aliena var. acuteserrata seeds and P. armandii seeds were transported to distant locations for feeding and burying. The Q. aliena var. acuteserrata seeds with diffusion distances of less than 1 m, 1—3 m and 5—8 m accounted for 59.12%, 18.23%, and 13.26% of predated seeds, respectively; the farthest diffusion distance was 12 m. The seed diffusion distance of P. armandii was mainly between 1—3 m(37.85%) and 3—5 m(23.73%), and was within the range of 5—8 m(13.56%), 8—10 m(11.86%), and greater than 10 m(11.30%);the maximum diffusion distance of P. armandii seeds was also 12 m. Rodents tended to transport Q. aliena var. acuteserrata seeds to the pine forest for feeding, while transporting the seeds of P. armandii to the oak forest, indicating that different forest types are an important factor in seed dispersal. P. tabuliformis seeds were the most predated on by rodents(96.90%), followed by Q. aliena var. acuteserrata seeds(73.57%), and P. armandii seeds(50%). In the following year′s survey, most of the predated seeds had been consumed, and only a handful of Q. aliena var. acuteserrata seeds had sprouted into seedlings(1.67%). Rodent predation and storage behaviors are critical for the spread of forest seeds and their successful renewal.
Rodents in forest communities have a feeding preference for tree seeds. This preference will lead to different predation and storage strategies for rodents, resulting in different diffusion patterns of tree seeds. In this study, the seed fate of Quercus aliena var. acuteserrata, Pinus armandii, and Pinus tabuliformis was studied to analyze the seed dispersal patterns of constructive species in the pine-oak mixed forests of the Qinling Mountains, Shaanxi Province, China. The results demonstrated that seed traits are the main factor affecting their removal by rodents. The in-situ seed predation rate of P. tabuliformis seeds was 83.33%, which was notably higher than that of other seed types, indicating that rodents preferentially fed on the small P. tabuliformis seeds. Q. aliena var. acuteserrata seeds and P. armandii seeds were transported to distant locations for feeding and burying. The Q. aliena var. acuteserrata seeds with diffusion distances of less than 1 m, 1—3 m and 5—8 m accounted for 59.12%, 18.23%, and 13.26% of predated seeds, respectively; the farthest diffusion distance was 12 m. The seed diffusion distance of P. armandii was mainly between 1—3 m(37.85%) and 3—5 m(23.73%), and was within the range of 5—8 m(13.56%), 8—10 m(11.86%), and greater than 10 m(11.30%);the maximum diffusion distance of P. armandii seeds was also 12 m. Rodents tended to transport Q. aliena var. acuteserrata seeds to the pine forest for feeding, while transporting the seeds of P. armandii to the oak forest, indicating that different forest types are an important factor in seed dispersal. P. tabuliformis seeds were the most predated on by rodents(96.90%), followed by Q. aliena var. acuteserrata seeds(73.57%), and P. armandii seeds(50%). In the following year′s survey, most of the predated seeds had been consumed, and only a handful of Q. aliena var. acuteserrata seeds had sprouted into seedlings(1.67%). Rodent predation and storage behaviors are critical for the spread of forest seeds and their successful renewal.
引文
[1] 康冰,王得祥,崔宏安,迪玮峙,杜焰玲.秦岭山地油松群落更新特征及影响因子.应用生态学报,2011,22(7):1659- 1667.
[2] 于飞.秦岭松栎混交林建群种种子扩散过程及天然更新格局研究[D].杨凌:西北农林科技大学,2014.
[3] Gómez J M,Puerta-Piňero C,Schupp E W.Effectiveness of rodents as local seed dispersers of Holm oaks.Oecologia,2008,155(3):529- 537.
[4] 张博,石子俊,陈晓宁,侯祥,王京,李金钢,常罡.森林鼠类对秦岭南坡3种壳斗科植物种子扩散的影响.生态学报,2016,36(21):6750- 6757.
[5] Chang G,Zhang Z B.Differences in hoarding behaviors among six sympatric rodent species on seeds of oil tea (Camellia oleifera) in Southwest China.Acta Oecologica,2011,37(3):165- 169.
[6] 肖治术,张知彬.啮齿动物的贮藏行为与植物种子的扩散.兽类学报,2004,24(1):61- 70.
[7] Yu F,Wang D X,Yi X F,Shi X X,Huang Y K,Zhang H W,Zhang X P.Does animal-mediated seed dispersal facilitate the formation of Pinus armandii-Quercus aliena var.acuteserrata forests?PLoS One,2014,9(2):e89886.
[8] Rusch U D,Midgley J J,Anderson B.Rodent consumption and caching behaviour selects for specific seed traits.South African Journal of Botany,2013,84:83- 87.
[9] 李宏俊,张知彬.动物与植物种子更新的关系Ⅱ.动物对种子的捕食、扩散、贮藏及与幼苗建成的关系.生物多样性,2001,9(1):25- 37.
[10] 陈晓宁,张博,陈雅娟,侯祥,王京,常罡.秦岭南北坡森林鼠类对板栗和锐齿栎种子扩散的影响.生态学报,2016,36(5):1303- 1311.
[11] 常罡,肖治术,张知彬.种子大小对小泡巨鼠贮藏行为的影响.兽类学报,2008,28(1):37- 41.
[12] 常罡,邰发道.季节变化对锐齿栎种子扩散的影响.生态学杂志,2011,30(1):189- 192.
[13] 周立彪,闫兴富,王建礼,周云峰.啮齿动物对不同大小和种皮特征种子的取食和搬运.应用生态学报,2013,24(8):2325- 2332.
[14] 陈晓宁,张博,王京,侯祥,韩宁,申健,常罡.食物源距离对中华姬鼠贮藏策略的影响.兽类学报,2017,37(2):146- 151.
[15] 于飞,牛可坤,焦广强,吕浩秋,易现峰.小型啮齿动物对小兴安岭5种林木种子扩散的影响.东北林业大学学报,2011,39(1):11- 13.
[16] 于飞,史晓晓,易现峰,王得祥.蒙古栎种子相对丰富度对小兴安岭5种木本植物种子扩散的影响.应用生态学报,2013,24(6):1531- 1535.
[17] 刘国强,刘长渠,易现峰.胡桃楸种子大小对鼠类分散贮藏行为的影响——基于无线电标记技术.生态学报,2015,35(17):5648- 5653.
[18] 欧瑞东,闫兴富,马惠娟,姜英娜.啮齿动物对不同大小辽东栎和山桃种子的取食和搬运.种子,2017,36(9):76- 80.
[19] Yu F,Shi X X,Wang D X,Yi X F,Fan D F,Guo T D,Lou Y G.Effects of insect infestation on Quercus aliena var.acuteserrata acorn dispersal in the Qinling Mountains,China.New Forests,2015,46(1):51- 61.
[20] 康海斌,王得祥,常明捷,康冰,胡有宁,于飞,庞越,滕青林.啮齿动物对不同林木种子的搬运和取食微生境选择机制.生态学报,2017,37(22):7604- 7613.
[21] Xiao Z S,Jansen P A,Zhang Z B.Using seed-tagging methods for assessing post-dispersal seed fate in rodent-dispersed trees.Forest Ecology and Management,2006,223(1/3):18- 23.
[22] 常罡.鼠类扩散种子的几种标签标记法的比较.生态学杂志,2012,31(3):684- 688.
[23] Perea R,Miguel A S,Gil L.Flying vs.climbing:factors controlling arboreal seed removal in oak-beech forests.Forest Ecology and Management,2011,262(7):1251- 1257.
[24] 沈泽昊,唐圆圆,李道兴.啮齿类取食的物种偏好与时空格局.生态学报,2008,28(12):6018- 6024.
[25] 王冲,张义锋,王振龙,乔王铁,路纪琪.济源太行山区鼠类对三种林木种子的扩散和贮藏.兽类学报,2013,33(2):150- 156.
[26] Lei J J,Shen Z,Yi X F.Pericarp thickness and seed size determine acorn dispersal of five rodent-dispersed oak species.Acta Theriologica Sinica,2012,32(2):83- 89.
[27] Chang G,Jin T Z,Pei J F,Chen X N,Zhang B,Shi Z J.Seed dispersal of three sympatric oak species by forest rodents in the Qinling Mountains,Central China.Plant Ecology,2012,213(10):1633- 1642.
[28] Zhang H M,Zhang Z B.Endocarp thickness affects seed removal speed by small rodents in a warm-temperate broad-leafed deciduous forest,China.Acta Oecologica,2008,34(3):285- 293.
[29] 路纪琪,张知彬.捕食风险及其对动物觅食行为的影响.生态学杂志,2004,23(2):66- 72.
[30] Dally J M,Clayton N S,Emery N J.The behaviour and evolution of cache protection and pilferage.Animal Behaviour,2006,72(1):13- 23.
[31] Vieira E M,Pizo M A,Izar P.Fruit and seed exploitation by small rodents of the Brazilian Atlantic forest.Mammalia,2003,67(4):533- 540.
[32] Gómez J M.Spatial patterns in long-distance dispersal of Quercus ilex acorns by jays in a heterogeneous landscape.Ecography,2003,26(5):573- 584.
[33] 张洪武,于飞,王得祥,张子良.秦岭南坡松栎林群落演替过程中物种组成及多样性动态分析.西北植物学报,2014,34(1):169- 176.
[34] 闫兴富,周立彪,刘建利.啮齿动物捕食压力下生境类型和覆盖处理对辽东栎种子命运的影响.生态学报,2012,32(9):2778- 2787.
[35] 路纪琪,张知彬.灌丛高度对啮齿动物贮藏和扩散辽东栎坚果的影响.动物学报,2005,51(2):195- 204.
[36] Cao L,Xiao Z S,Guo C,Chen J.Scatter-hoarding rodents as secondary seed dispersers of a frugivore-dispersed tree Scleropyrum wallichianum in a defaunated Xishuangbanna tropical forest,China.Integrative Zoology,2011,6(3):227- 234.
[37] 孙明洋,马庆亮,田澍辽,王建东,路纪琪.种子产量对鼠类扩散栓皮栎坚果的影响.兽类学报,2011,31(3):265- 271.
[38] 张树梓,李梅,张树彬,张志东,黄选瑞.塞罕坝华北落叶松人工林天然更新影响因子.生态学报,2015,35(16):5403- 5411.
[39] 李登武,王冬梅,姚文旭.油松的自毒作用及其生态学意义.林业科学,2010,46(11):174- 178.
[2] 于飞.秦岭松栎混交林建群种种子扩散过程及天然更新格局研究[D].杨凌:西北农林科技大学,2014.
[3] Gómez J M,Puerta-Piňero C,Schupp E W.Effectiveness of rodents as local seed dispersers of Holm oaks.Oecologia,2008,155(3):529- 537.
[4] 张博,石子俊,陈晓宁,侯祥,王京,李金钢,常罡.森林鼠类对秦岭南坡3种壳斗科植物种子扩散的影响.生态学报,2016,36(21):6750- 6757.
[5] Chang G,Zhang Z B.Differences in hoarding behaviors among six sympatric rodent species on seeds of oil tea (Camellia oleifera) in Southwest China.Acta Oecologica,2011,37(3):165- 169.
[6] 肖治术,张知彬.啮齿动物的贮藏行为与植物种子的扩散.兽类学报,2004,24(1):61- 70.
[7] Yu F,Wang D X,Yi X F,Shi X X,Huang Y K,Zhang H W,Zhang X P.Does animal-mediated seed dispersal facilitate the formation of Pinus armandii-Quercus aliena var.acuteserrata forests?PLoS One,2014,9(2):e89886.
[8] Rusch U D,Midgley J J,Anderson B.Rodent consumption and caching behaviour selects for specific seed traits.South African Journal of Botany,2013,84:83- 87.
[9] 李宏俊,张知彬.动物与植物种子更新的关系Ⅱ.动物对种子的捕食、扩散、贮藏及与幼苗建成的关系.生物多样性,2001,9(1):25- 37.
[10] 陈晓宁,张博,陈雅娟,侯祥,王京,常罡.秦岭南北坡森林鼠类对板栗和锐齿栎种子扩散的影响.生态学报,2016,36(5):1303- 1311.
[11] 常罡,肖治术,张知彬.种子大小对小泡巨鼠贮藏行为的影响.兽类学报,2008,28(1):37- 41.
[12] 常罡,邰发道.季节变化对锐齿栎种子扩散的影响.生态学杂志,2011,30(1):189- 192.
[13] 周立彪,闫兴富,王建礼,周云峰.啮齿动物对不同大小和种皮特征种子的取食和搬运.应用生态学报,2013,24(8):2325- 2332.
[14] 陈晓宁,张博,王京,侯祥,韩宁,申健,常罡.食物源距离对中华姬鼠贮藏策略的影响.兽类学报,2017,37(2):146- 151.
[15] 于飞,牛可坤,焦广强,吕浩秋,易现峰.小型啮齿动物对小兴安岭5种林木种子扩散的影响.东北林业大学学报,2011,39(1):11- 13.
[16] 于飞,史晓晓,易现峰,王得祥.蒙古栎种子相对丰富度对小兴安岭5种木本植物种子扩散的影响.应用生态学报,2013,24(6):1531- 1535.
[17] 刘国强,刘长渠,易现峰.胡桃楸种子大小对鼠类分散贮藏行为的影响——基于无线电标记技术.生态学报,2015,35(17):5648- 5653.
[18] 欧瑞东,闫兴富,马惠娟,姜英娜.啮齿动物对不同大小辽东栎和山桃种子的取食和搬运.种子,2017,36(9):76- 80.
[19] Yu F,Shi X X,Wang D X,Yi X F,Fan D F,Guo T D,Lou Y G.Effects of insect infestation on Quercus aliena var.acuteserrata acorn dispersal in the Qinling Mountains,China.New Forests,2015,46(1):51- 61.
[20] 康海斌,王得祥,常明捷,康冰,胡有宁,于飞,庞越,滕青林.啮齿动物对不同林木种子的搬运和取食微生境选择机制.生态学报,2017,37(22):7604- 7613.
[21] Xiao Z S,Jansen P A,Zhang Z B.Using seed-tagging methods for assessing post-dispersal seed fate in rodent-dispersed trees.Forest Ecology and Management,2006,223(1/3):18- 23.
[22] 常罡.鼠类扩散种子的几种标签标记法的比较.生态学杂志,2012,31(3):684- 688.
[23] Perea R,Miguel A S,Gil L.Flying vs.climbing:factors controlling arboreal seed removal in oak-beech forests.Forest Ecology and Management,2011,262(7):1251- 1257.
[24] 沈泽昊,唐圆圆,李道兴.啮齿类取食的物种偏好与时空格局.生态学报,2008,28(12):6018- 6024.
[25] 王冲,张义锋,王振龙,乔王铁,路纪琪.济源太行山区鼠类对三种林木种子的扩散和贮藏.兽类学报,2013,33(2):150- 156.
[26] Lei J J,Shen Z,Yi X F.Pericarp thickness and seed size determine acorn dispersal of five rodent-dispersed oak species.Acta Theriologica Sinica,2012,32(2):83- 89.
[27] Chang G,Jin T Z,Pei J F,Chen X N,Zhang B,Shi Z J.Seed dispersal of three sympatric oak species by forest rodents in the Qinling Mountains,Central China.Plant Ecology,2012,213(10):1633- 1642.
[28] Zhang H M,Zhang Z B.Endocarp thickness affects seed removal speed by small rodents in a warm-temperate broad-leafed deciduous forest,China.Acta Oecologica,2008,34(3):285- 293.
[29] 路纪琪,张知彬.捕食风险及其对动物觅食行为的影响.生态学杂志,2004,23(2):66- 72.
[30] Dally J M,Clayton N S,Emery N J.The behaviour and evolution of cache protection and pilferage.Animal Behaviour,2006,72(1):13- 23.
[31] Vieira E M,Pizo M A,Izar P.Fruit and seed exploitation by small rodents of the Brazilian Atlantic forest.Mammalia,2003,67(4):533- 540.
[32] Gómez J M.Spatial patterns in long-distance dispersal of Quercus ilex acorns by jays in a heterogeneous landscape.Ecography,2003,26(5):573- 584.
[33] 张洪武,于飞,王得祥,张子良.秦岭南坡松栎林群落演替过程中物种组成及多样性动态分析.西北植物学报,2014,34(1):169- 176.
[34] 闫兴富,周立彪,刘建利.啮齿动物捕食压力下生境类型和覆盖处理对辽东栎种子命运的影响.生态学报,2012,32(9):2778- 2787.
[35] 路纪琪,张知彬.灌丛高度对啮齿动物贮藏和扩散辽东栎坚果的影响.动物学报,2005,51(2):195- 204.
[36] Cao L,Xiao Z S,Guo C,Chen J.Scatter-hoarding rodents as secondary seed dispersers of a frugivore-dispersed tree Scleropyrum wallichianum in a defaunated Xishuangbanna tropical forest,China.Integrative Zoology,2011,6(3):227- 234.
[37] 孙明洋,马庆亮,田澍辽,王建东,路纪琪.种子产量对鼠类扩散栓皮栎坚果的影响.兽类学报,2011,31(3):265- 271.
[38] 张树梓,李梅,张树彬,张志东,黄选瑞.塞罕坝华北落叶松人工林天然更新影响因子.生态学报,2015,35(16):5403- 5411.
[39] 李登武,王冬梅,姚文旭.油松的自毒作用及其生态学意义.林业科学,2010,46(11):174- 178.