荒漠孑遗植物裸果木种子时空扩散特性
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摘要
裸果木(Gymnocarpos przewalskii)是亚洲中部荒漠区少有的第三纪孑遗物种,由于气候变化及人为干扰,其自然种群分布范围不断缩小。种子扩散作为植物生活史过程中的重要阶段,不仅对物种生存及其多样性至关重要,还影响物种分布范围和局部丰度。2015年和2016年分别在新疆哈密地区,采用布设种子收集器的方法,对其自然种群种子扩散的时空动态进行了定点连续观测。结果表明:该物种于当年6月上旬开始扩散,2015年略早于2016年。每年种子扩散持续时间约两个月,扩散趋势为单峰曲线,且呈集中大量扩散的模式,扩散高峰期与当年初次月降水高峰期吻合;在顺风的正南和东南方向上,种子扩散密度大且距离远;种子扩散主要集中在母株冠幅下,随着距母株距离的增加,种子扩散密度减少,二者间存在极显著的负相关性(P<0.01),由于裸果木枝条繁多,对风力强度起到了一定的阻碍作用,可能是造成种子集中扩散在母株下的原因。裸果木种子扩散受外界环境(降水、风向)和自身因素等方面的影响,当种子在大量降水前完成扩散,将有利于种子在适宜的微生境萌发,是对多风、干旱的恶劣生境的一种长期适应。
Gymnocarpos przewalskii is a rare Tertiary relict species found in the desert regions of Central Asia; the natural population distribution of G. przewalskii is declining owing to climate change and human interference. As an important life history stage of plants, seed dispersal is not only critical for the survival and diversity of species, but also affects the distribution and local abundance of species. In 2015 and 2016, seed traps were installed around G. przewalskii mother plants in different directions in Hami. The spatial and temporal dynamics of seed dispersal of this natural population were continuously observed. The results showed that seeds began to disperse in early June and seed dispersal in 2015 was slightly earlier than in 2016. Seed dispersal lasted about 2 months; the dispersal trend was a single peak curve and presents a pattern of mass dispersal. The peak of seed dispersal coincided with the peak of the first monthly precipitation of the year. Dispersal occurred at the highest density and for the longest distance south and southeast of downwind. The seeds were mainly concentrated directly underneath the canopy of the mother plant. As the distance from the mother plant increased, seed density decreased, and there was a very significant negative correlation between dispersal distance and dispersal density. The main reason for the high dispersal density of seeds of G. przewalskii under the mother plant is that its numerous branches weakened the wind strength through the plants. Seed dispersal of G. przewalskii was affected by precipitation, wind, and plant characteristics. Seed dispersal before heavy precipitation may be beneficial for seed germination in suitable microhabitats, which is a long-term adaptation of G. przewalskii to the harsh wind and drought conditions.
Gymnocarpos przewalskii is a rare Tertiary relict species found in the desert regions of Central Asia; the natural population distribution of G. przewalskii is declining owing to climate change and human interference. As an important life history stage of plants, seed dispersal is not only critical for the survival and diversity of species, but also affects the distribution and local abundance of species. In 2015 and 2016, seed traps were installed around G. przewalskii mother plants in different directions in Hami. The spatial and temporal dynamics of seed dispersal of this natural population were continuously observed. The results showed that seeds began to disperse in early June and seed dispersal in 2015 was slightly earlier than in 2016. Seed dispersal lasted about 2 months; the dispersal trend was a single peak curve and presents a pattern of mass dispersal. The peak of seed dispersal coincided with the peak of the first monthly precipitation of the year. Dispersal occurred at the highest density and for the longest distance south and southeast of downwind. The seeds were mainly concentrated directly underneath the canopy of the mother plant. As the distance from the mother plant increased, seed density decreased, and there was a very significant negative correlation between dispersal distance and dispersal density. The main reason for the high dispersal density of seeds of G. przewalskii under the mother plant is that its numerous branches weakened the wind strength through the plants. Seed dispersal of G. przewalskii was affected by precipitation, wind, and plant characteristics. Seed dispersal before heavy precipitation may be beneficial for seed germination in suitable microhabitats, which is a long-term adaptation of G. przewalskii to the harsh wind and drought conditions.
引文
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[2] Nathan R,Schurr F M,Spiegel O,Steinitz O,Trakhtenbrot A,Soar A.Mechanisms of long-distance seed dispersal.Trends in Ecology & Evolution,2008,23(11):638- 647.
[3] Chanyenga T F,Geldenhuys C J,Harvey J.Variation in seed rain from Widdringtonia whytei growing in different conditions on Mulanje Mountain in Malawi.Southern Forests:A Journal of Forest Science,2011,73(3/4):123- 129.
[4] 朱金雷,刘志民.种子传播生物学主要术语和概念.生态学杂志,2012,31(9):2397- 2403.
[5] Gutterman Y.Strategies of seed dispersal and germination in plants inhabiting deserts.The Botanical Review,1994,60(4):373- 425.
[6] Howe H F,Smallwood J.Ecology of seed dispersal.Annual Review of Ecology and Systematics,1982,13:201- 228.
[7] Wang B C,Smith T B.Closing the seed dispersal loop.Trends in Ecology & Evolution,2002,17(8):379- 386.
[8] Bullock J M,Shea K,Skarpaas O.Measuring plant dispersal:an introduction to field methods and experimental design.Plant Ecology,2006,186(2):217- 234.
[9] Clark J S,Silman M,Kern R,Macklin E,Hillerislambers J.Seed dispersal near and far:patterns across temperate and tropical forests.Ecology,1999,80(5):1475- 1494.
[10] Howe H F,Miriti M N.No question:seed dispersal matters.Trends in Ecology & Evolution,2000,15(11):434- 436.
[11] 胡星明,蔡永立,李恺,张天澍.浙江天童常绿阔叶林栲树种子雨的时空格局.应用生态学报,2005,16(5):815- 819.
[12] 刘双,金光泽.小兴安岭阔叶红松(Pinus koraiensis)林种子雨的时空动态.生态学报,2008,28(11):5731- 5740.
[13] 谷伟,岳永杰,李钢铁,潘文杰,哈丽雅.浑善达克沙地沙地榆种子雨的扩散规律.生态学报,2012,32(11):3440- 3448.
[14] Liu H L,Zhang D Y,Yang X J,Huang Z Y,Duan S M,Wang X Y.Seed dispersal and germination traits of 70 plant species inhabiting the Gurbantunggut desert in northwest China.The Scientific World Journal,2014,2014:346405.
[15] 王进,颜霞,李军元,张勇,谢全刚,李刚,顾文豪.蒙古扁桃种子雨散布及种子萌发特性研究.冰川冻土,2017,39(2):429- 434.
[16] Zhao X Y,Baskin C C,Zhu C Q,Baskin J M.Fruit dispersal dynamics of the cold desert shrub Zygophyllum xanthoxylon.Seed Science Research,2017,27(4):303- 310.
[17] 柴永青,曹致中,蔡卓山,万丽霞,李学玲.肃北地区稀有植物裸果木种群的空间分布格局.草业学报,2010,19(5):239- 249.
[18] 赵哈林.沙漠化过程中植物的适应对策及植被稳定性机理.北京:海洋出版社,2004:91- 96.
[19] Trakhtenbrot A,Katul G G,Nathan R.Mechanistic modeling of seed dispersal by wind over hilly terrain.Ecological Modelling,2014,274:29- 40.
[20] 王立龙,王亮,张丽芳,刘玉洋,徐世健.不同生境下濒危植物裸果木种群结构及动态特征.植物生态学报,2015,39(10):980- 989.
[21] 王博,王亮,王立龙,杨增武,姚静斐,徐世健.濒危植物裸果木(Gymnocarpos przewalskii)与其伴生种种间联结性及群落稳定性.中国沙漠,2017,37(1):86- 92.
[22] 李新蓉,唐欣,付文洁.荒漠孑遗植物裸果木的花部特征及繁育系统.生态学杂志,2016,35(10):2592- 2598.
[23] 唐欣,李新蓉.荒漠孑遗植物裸果木的开花物候特征.植物学报,2017,52(4):487- 495.
[24] 杨赵平,梁继业,柴学平,薛婉婷.环境因子对珍稀濒危植物裸果木种子萌发特性的影响.西南林业大学学报,2017,37(1):49- 53.
[25] 张玉波,李景文,张昊,邹大林,武逢平,程春龙,李俊清,李帅英.胡杨种子散布的时空分布格局.生态学报,2005,25(8):1994- 2000.
[26] 龙丽红,王慧,马晓丽,徐鸿斌,王绍明.古尔班通古特沙漠羽毛针禾(Stipagrostis pennata)种群种子雨特征.干旱区研究,2014,31(3):516- 522.
[27] 吕朝燕,张希明,刘国军,吴俊侠,闫海龙,邓潮洲.准噶尔盆地东南缘梭梭种子雨特征.生态学报,2012,32(19):6270- 6278.
[28] Culot L,Muňoz Lazo F J J,Huynen M C,Poncin P,Heymann E W.Seasonal variation in seed dispersal by tamarins alters seed rain in a secondary rain forest.International Journal of Primatology,2010,31(4):553- 569.
[29] 李娟,郭聪,肖治术.都江堰亚热带森林常见木本植物果实组成与种子扩散策略.生物多样性,2013,21(5):572- 581.
[30] 查同刚,孙向阳,王登芝,阎海平.北京西山地区人工侧柏林种子雨的研究.北京林业大学学报,2003,25(1):28- 31.
[31] Ramos D M,Diniz P,Ooi M K J,Borghetti F,Valls J F M.Avoiding the dry season:dispersal time and syndrome mediate seed dormancy in grasses in Neotropical savanna and wet grasslands.Journal of Vegetation Science,2017,28(4):798- 807.
[32] Heydel F,Cunze S,Bernhardt-R?mermann M,Tackenberg O.Long-distance seed dispersal by wind:disentangling the effects of species traits,vegetation types,vertical turbulence and wind speed.Ecological Research,2014,29(4):641- 651.
[33] 于顺利,郎南军,彭明俊,赵琳,郭永清,郑科,张立新,温绍龙,李晖.种子雨研究进展.生态学杂志,2007,26(10):1646- 1652.
[34] 赵丽娅,李兆华,赵锦慧,赵哈林,李锋瑞.流动沙丘先锋灌木差巴嘎蒿的种子散布格局.西北植物学报,2005,25(9):1821- 1826.
[35] 白成科,曹博,李桂双.植物种子传播途径与基因组值和千粒重的相关性.生态学杂志,2013,32(4):832- 837.
[2] Nathan R,Schurr F M,Spiegel O,Steinitz O,Trakhtenbrot A,Soar A.Mechanisms of long-distance seed dispersal.Trends in Ecology & Evolution,2008,23(11):638- 647.
[3] Chanyenga T F,Geldenhuys C J,Harvey J.Variation in seed rain from Widdringtonia whytei growing in different conditions on Mulanje Mountain in Malawi.Southern Forests:A Journal of Forest Science,2011,73(3/4):123- 129.
[4] 朱金雷,刘志民.种子传播生物学主要术语和概念.生态学杂志,2012,31(9):2397- 2403.
[5] Gutterman Y.Strategies of seed dispersal and germination in plants inhabiting deserts.The Botanical Review,1994,60(4):373- 425.
[6] Howe H F,Smallwood J.Ecology of seed dispersal.Annual Review of Ecology and Systematics,1982,13:201- 228.
[7] Wang B C,Smith T B.Closing the seed dispersal loop.Trends in Ecology & Evolution,2002,17(8):379- 386.
[8] Bullock J M,Shea K,Skarpaas O.Measuring plant dispersal:an introduction to field methods and experimental design.Plant Ecology,2006,186(2):217- 234.
[9] Clark J S,Silman M,Kern R,Macklin E,Hillerislambers J.Seed dispersal near and far:patterns across temperate and tropical forests.Ecology,1999,80(5):1475- 1494.
[10] Howe H F,Miriti M N.No question:seed dispersal matters.Trends in Ecology & Evolution,2000,15(11):434- 436.
[11] 胡星明,蔡永立,李恺,张天澍.浙江天童常绿阔叶林栲树种子雨的时空格局.应用生态学报,2005,16(5):815- 819.
[12] 刘双,金光泽.小兴安岭阔叶红松(Pinus koraiensis)林种子雨的时空动态.生态学报,2008,28(11):5731- 5740.
[13] 谷伟,岳永杰,李钢铁,潘文杰,哈丽雅.浑善达克沙地沙地榆种子雨的扩散规律.生态学报,2012,32(11):3440- 3448.
[14] Liu H L,Zhang D Y,Yang X J,Huang Z Y,Duan S M,Wang X Y.Seed dispersal and germination traits of 70 plant species inhabiting the Gurbantunggut desert in northwest China.The Scientific World Journal,2014,2014:346405.
[15] 王进,颜霞,李军元,张勇,谢全刚,李刚,顾文豪.蒙古扁桃种子雨散布及种子萌发特性研究.冰川冻土,2017,39(2):429- 434.
[16] Zhao X Y,Baskin C C,Zhu C Q,Baskin J M.Fruit dispersal dynamics of the cold desert shrub Zygophyllum xanthoxylon.Seed Science Research,2017,27(4):303- 310.
[17] 柴永青,曹致中,蔡卓山,万丽霞,李学玲.肃北地区稀有植物裸果木种群的空间分布格局.草业学报,2010,19(5):239- 249.
[18] 赵哈林.沙漠化过程中植物的适应对策及植被稳定性机理.北京:海洋出版社,2004:91- 96.
[19] Trakhtenbrot A,Katul G G,Nathan R.Mechanistic modeling of seed dispersal by wind over hilly terrain.Ecological Modelling,2014,274:29- 40.
[20] 王立龙,王亮,张丽芳,刘玉洋,徐世健.不同生境下濒危植物裸果木种群结构及动态特征.植物生态学报,2015,39(10):980- 989.
[21] 王博,王亮,王立龙,杨增武,姚静斐,徐世健.濒危植物裸果木(Gymnocarpos przewalskii)与其伴生种种间联结性及群落稳定性.中国沙漠,2017,37(1):86- 92.
[22] 李新蓉,唐欣,付文洁.荒漠孑遗植物裸果木的花部特征及繁育系统.生态学杂志,2016,35(10):2592- 2598.
[23] 唐欣,李新蓉.荒漠孑遗植物裸果木的开花物候特征.植物学报,2017,52(4):487- 495.
[24] 杨赵平,梁继业,柴学平,薛婉婷.环境因子对珍稀濒危植物裸果木种子萌发特性的影响.西南林业大学学报,2017,37(1):49- 53.
[25] 张玉波,李景文,张昊,邹大林,武逢平,程春龙,李俊清,李帅英.胡杨种子散布的时空分布格局.生态学报,2005,25(8):1994- 2000.
[26] 龙丽红,王慧,马晓丽,徐鸿斌,王绍明.古尔班通古特沙漠羽毛针禾(Stipagrostis pennata)种群种子雨特征.干旱区研究,2014,31(3):516- 522.
[27] 吕朝燕,张希明,刘国军,吴俊侠,闫海龙,邓潮洲.准噶尔盆地东南缘梭梭种子雨特征.生态学报,2012,32(19):6270- 6278.
[28] Culot L,Muňoz Lazo F J J,Huynen M C,Poncin P,Heymann E W.Seasonal variation in seed dispersal by tamarins alters seed rain in a secondary rain forest.International Journal of Primatology,2010,31(4):553- 569.
[29] 李娟,郭聪,肖治术.都江堰亚热带森林常见木本植物果实组成与种子扩散策略.生物多样性,2013,21(5):572- 581.
[30] 查同刚,孙向阳,王登芝,阎海平.北京西山地区人工侧柏林种子雨的研究.北京林业大学学报,2003,25(1):28- 31.
[31] Ramos D M,Diniz P,Ooi M K J,Borghetti F,Valls J F M.Avoiding the dry season:dispersal time and syndrome mediate seed dormancy in grasses in Neotropical savanna and wet grasslands.Journal of Vegetation Science,2017,28(4):798- 807.
[32] Heydel F,Cunze S,Bernhardt-R?mermann M,Tackenberg O.Long-distance seed dispersal by wind:disentangling the effects of species traits,vegetation types,vertical turbulence and wind speed.Ecological Research,2014,29(4):641- 651.
[33] 于顺利,郎南军,彭明俊,赵琳,郭永清,郑科,张立新,温绍龙,李晖.种子雨研究进展.生态学杂志,2007,26(10):1646- 1652.
[34] 赵丽娅,李兆华,赵锦慧,赵哈林,李锋瑞.流动沙丘先锋灌木差巴嘎蒿的种子散布格局.西北植物学报,2005,25(9):1821- 1826.
[35] 白成科,曹博,李桂双.植物种子传播途径与基因组值和千粒重的相关性.生态学杂志,2013,32(4):832- 837.
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