天山雪岭云杉森林群落的密度制约效应

doi:10.17520/biods.2015282

摘要/Abstract

摘要：

密度制约是维持自然森林群落物种共存的重要机制之一。雪岭云杉(Picea schrenkiana)是天山森林群落的单优种, 在我国温带森林中占有重要地位。本文基于对天山雪岭云杉8 ha森林动态监测样地的两次调查结果, 分析了该样地群落物种组成的动态变化, 运用点格局方法分析了雪岭云杉个体在不同龄级的空间分布格局及其关联性, 探讨了该群落的密度制约效应。结果表明: (1)该样地内, DBH ≥ 1 cm的木本植物在2009年有11,835株, 2014年为11,050株, 5年间的个体死亡率为8.82%, 补员率为2.19%, 个体株数净减少6.63%。(2)使用双关联函数g(r)分析了不同龄级雪岭云杉的空间分布格局, 发现幼龄树和中龄树在0-40 m尺度上主要呈现聚集分布, 但随着尺度增大聚集强度逐渐减小; 老龄树在0-2 m及4 m尺度上呈随机分布, 在其他尺度上呈聚集分布, 但聚集程度较小。(3)用“案例-对照”设计的方法, 排除生境异质性的影响后, 将幼龄树和中龄树作为案例, 老龄树作为对照, 对比幼龄树、中龄树与老龄树的分布格局, 发现雪岭云杉幼龄树和中龄树在0-40 m尺度上呈现额外的聚集, 随着径级的增大, 这种额外的聚集强度逐渐减小, 即表现出密度制约效应。(4)老龄树与中龄树、幼龄树的空间关联性相同, 在0-40 m尺度上均呈明显负关联。

关键词: 动态监测样地, 密度制约假说, 空间分布, 雪岭云杉, 天山

Abstract

The density-dependence hypothesis is one of the most important mechanisms proposed to explain species coexistence in natural forest communities. Picea schrenkiana is the single dominant species of the forest communities in Tianshan Mountains, which plays a very important role in temperate forests of China. We used reexamined data with a five-year interval on an 8-ha P. schrenkiana forest dynamics plot to illustrate the dynamics of species composition. Pointpattern analysis was conducted to reveal the spatial structure of P. schrenkiana individuals of different ages, and the effects of the density-dependence hypothesis on different ages of P. schrenkiana was discussed. The results were as follows: (1) Woody plants with DBH ≥ 1 cm were 11,835 trees in 2009 and 11,050 in 2014. The mortality rate was 8.82%, the recruitment rate was 2.19%, and the net decrease was 6.63%; (2) Young and medium trees of P. schrenkiana presented an aggregation distribution across a scale of 0-40 m, and the aggregation intensity decreased as the scale increased. Mature trees presented a random distribution on scales of 0 m ≤ r ≤ 2 m and r = 4 m, and an aggregation distribution on other scales with a low density. (3) Using a “case-control” design method and eliminating habitat heterogeneity, the distribution of young and medium trees were taken as examples of case. The distribution pattern of mature trees was used as a comparison and represented habitat heterogeneity, and was compared with that of young and medium trees. Young and medium trees of P. schrenkiana were observed to present more considerable gathering than was observed for mature trees on scales of 0-40 m, and density decreased while the diameter class increased. These results indicate that the density-dependence hypothesis has an influence on distribution patterns of different age classes. This study reveals that P. schrenkiana was affected by habitat heterogeneity and presented a significant aggregation effect. The spatial pattern of P. schrenkiana is revealed to be affected by the density-dependence hypothesis after eliminating the effects of habitat heterogeneity. The results of this paper support the Janzen-Connell hypothesis.

Key words: dynamics plot, density-dependence hypothesis, spatial distribution, Picea schrenkiana, Tianshan Mountains

王慧杰, 常顺利, 张毓涛, 谢锦, 何平, 宋成程, 孙雪娇 (2016) 天山雪岭云杉森林群落的密度制约效应. 生物多样性, 24, 252-261. DOI: 10.17520/biods.2015282.

Huijie Wang, Shunli Chang, Yutao Zhang, Jin Xie, Ping He, Chengcheng Song, Xuejiao Sun (2016) Density-dependent effects in Picea schrenkiana forests in Tianshan Mountains. Biodiversity Science, 24, 252-261. DOI: 10.17520/biods.2015282.

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链接本文: https://www.biodiversity-science.net/CN/10.17520/biods.2015282

https://www.biodiversity-science.net/CN/Y2016/V24/I3/252

图/表 6

图1 天山雪岭云杉8 ha森林动态监测样地示意图

Fig. 1 The sketch map of the 8-ha Picea schrenkiana forest dynamics plot in Tianshan Mountains

表1 2009和2014年天山雪岭云杉8 ha森林动态监测样地的群落动态

Table 1 Community dynamics in the 8-ha Picea schrenkiana forest dynamics plot in Tianshan Mountains in 2009 and 2014

物种 Species	生长型 Growth form	多度 Abundance			重要值 Importance value (%)		平均胸径 Mean DBH (cm)			死亡数 Number of dead individual	补员数 Number of recruited individual	胸径净增长 DBH net growth (cm)
物种 Species	生长型 Growth form	2009	2014		2009	2014		2009	2014	死亡数 Number of dead individual	补员数 Number of recruited individual	胸径净增长 DBH net growth (cm)
雪岭云杉 Picea schrenkiana	乔木 Tree	11,731	10,962	92.88		92.51	13.16		14.25	1,010	241	1.03
密叶杨 Populus talassica	乔木 Tree	5	5	0.42		0.41	9.20		10.88	0	0	1.68
天山桦 Betula tianschanica	乔木 Tree	1	0	0.14		0.00	4.90		5.20	1	0	-
天山花楸 Sorbus tianschanica	乔木 Tree	31	25	1.45		2.58	8.31		7.81	8	2	0.68
小檗 Berberis heteropoda	灌木 Shrub	34	40	2.22		2.74	2.11		2.05	8	14	0.16
忍冬 Lonicera hispida	灌木 Shrub	27	17	2.08		1.63	4.91		5.94	11	1	0.80
栒子 Cotoneaster melanocarpus	灌木 Shrub	4	0	0.54		0.00	5.56		0.00	4	0	-
蔷薇 Rosa spinosissima	灌木 Shrub	2	1	0.27		0.13	4.60		3.30	2	1	-
合计 Total		11,835	11,050	100		100				1,044	259

图2 天山雪岭云杉8 ha森林动态监测样地内不同龄级(径级)雪岭云杉的空间分布格局。(A)幼龄树(1 cm ≤ DBH ≤ 12.5 cm); (B)中龄树(12.5 cm < DBH ≤ 22.5 cm); (C)老龄树(DBH > 22.5 cm)。

Fig. 2 Spatial distribution patterns of Picea schrenkiana with different ages (DBH) in the 8-ha forest dynamics plot in Tianshan Mountains. (A) Young tree (1 cm ≤ DBH ≤ 12.5 cm); (B) Medium tree (12.5 cm < DBH ≤ 22.5 cm); (C) Mature tree (DBH > 22.5 cm).

图3 天山雪岭云杉8 ha森林动态监测样地内不同龄级(径级)雪岭云杉个体空间分布格局的尺度依赖性。(A)幼龄树(1 cm ≤ DBH ≤ 12.5 cm); (B)中龄树(12.5 cm < DBH ≤ 22.5 cm); (C)老龄树(DBH > 22.5 cm)。实线为99%置信区间的包迹线, 点线为实际点格局。

Fig. 3 Scale dependency of spatial patterns of Picea schrenkiana with different ages (DBH) in the 8-ha forest dynamics plot in Tianshan Mountains. (A) Young trees (1 cm ≤ DBH ≤ 12.5 cm); (B) Medium trees (12.5 cm < DBH ≤ 22.5 cm); (C) Mature trees (DBH > 22.5 cm). Solid lines show 99% simulation envelopes and solid lines with points represent the point pattern.

图4 天山雪岭云杉8 ha森林动态监测样地密度制约效应分析。(A)幼龄树(1 cm ≤ DBH ≤ 12.5 cm)作为案例, 老龄树(DBH > 22.5 cm)作为对照; (B)中龄树(12.5 cm < DBH ≤ 22.5 cm)作为案例, 老龄树作为对照; (C)从幼龄树到中龄树聚集格局的变化。实线为99%置信区间的包迹线, 点线为实际点的格局。

Fig. 4 Density-dependence analyses of the 8-ha Picea schrenkiana forest dynamics plot in Tianshan Mountains. (A) Young trees (1 cm ≤ DBH ≤ 12.5 cm) serve as case and mature trees (DBH > 22.5 cm) as control; (B) Medium trees (12.5 cm < DBH ≤ 22.5 cm) serve as case and mature trees as control; (C) The pattern changes from young trees to medium trees. Solid lines show 99% simulation envelopes and solid lines with points represents the point pattern.

图5 天山雪岭云杉8 ha森林动态监测样地内不同龄级(径级)雪岭云杉的空间关联性。(A)幼龄树(1 cm ≤ DBH ≤ 12.5 cm)与老龄树(DBH > 22.5 cm); (B)中龄树(12.5 cm < DBH ≤ 22.5 cm)与老龄树。实线为99％置信区间的包迹线, 点线为实际点的格局。

Fig. 5 Spatial association of Picea schrenkiana with different ages (DBH) in the 8-ha forest dynamics plot in Tianshan Mountains. (A) Young (1 cm ≤ DBH ≤ 12.5 cm) and mature (DBH > 22.5 cm) trees; (B) Medium (12.5 cm < DBH ≤ 22.5 cm) and mature trees. Solid lines show 99% simulation envelopes and solid lines with points represents the point pattern.

参考文献 13

14	Guo YL, Lu ZJ, Wang QG, Lu JM, Xu YZ, Meng HJ, Liu HB, Zhang JX, Bao DC, Qiao XJ, Huang HD, Jiang MX (2015) Detecting density dependence from spatial patterns in a heterogeneous subtropical forest of Central China. Canadian Journal of Forest Research, 45, 710-720
15	Guo YL, Wang B, Xiang WS, Ding T, Lu SH, Huang FZ, Li DX, Wen SJ, He YL, Li XK (2015) Dynamics of density-dependent effects of tree species in a 15 ha seasonal rain forest plot in northern tropical karst in Nonggang, Guangxi, southern China. Chinese Science Bulletin, 60, 1602-1611.(in Chinese with English abstract)
	[郭屹立, 王斌, 向悟生,丁涛, 陆树华, 黄甫昭, 李冬兴, 文淑均, 何运林, 李先琨 (2015)弄岗喀斯特季节性雨林15 ha样地密度制约效应分析. 科学通报, 60, 1602-1611.]
16	Harms KE, Wright SJ, Calderon O, Hernandez A, Herre EA (2000) Pervasive density-dependent recruitment enhances seedling diversity in a tropical forest. Nature, 404, 493-495.
17	He F, Duncan RP (2000) Density-dependent effects on tree survival in an old-growth Douglas fir forest. Journal of Ecology, 88, 676-688.
18	Huston MA, DeAngelis DL (1994) Competition and coexistence: the effects of resource transport and supply rates. The American Naturalist, 144, 954-977.
19	Hubbell SP, Ahumada JA, Condit R, Foster RB (2001) Local neighborhood effects on long-term survival of individualtrees in a Neotropical forest. Ecological Research, 16, 859-875.
20	Janzen DH (1970) Herbivores and the number of tree species in tropical forests. The American Naturalist, 104, 501-528.
21	Li L, Chen JH, Ren HB, Mi XC, Yu MJ, Yang B (2010) Spatial patterns of Castanopsis eyrei and Schima superba in mid-subtropical broad-leaved evergreen forest in Gutianshan National Nature Reserve, China. Chinese Journal of Plant Ecology, 34, 241-252.(in Chinese with English abstract)
	[李立, 陈建华, 任海保, 米湘成, 于明坚, 杨波 (2010) 古田山常绿阔叶林优势树种甜槠和木荷的空间格局分析. 植物生态学报, 34, 241-252.]
22	Lin YC, Chang LW, Yang KC, Wang HH, Sun IF (2011) Point patterns of tree distribution determined by habitat heterogeneity and dispersal limitation. Oecologia, 165, 175-184.
23	Luo ZR (2011) The Effects of Density Dependence in Subtropical Evergreen Broad-leaved Forest of Baishanzu. PhD dissertation, Zhejiang University, Hangzhou.(in Chinese with English abstract)
	[骆争荣 (2011) 百山祖亚热带常绿阔叶林群落密度制约的作用. 博士学位论文, 浙江大学, 杭州.]
24	Luo ZR, Mi XC, Chen XR, Ye ZL, Ding BY (2012) Density dependence is not very prevalent in a heterogeneous subtropical forest. Oikos, 121, 1239-1250.
25	Ma KP (2013) Studies on biodiversity and ecosystem function via manipulation experiments. Biodiversity Science, 21 247-248.(in Chinese)
	[马克平 (2013) 生物多样性与生态系统功能的实验研究. 生物多样性, 21, 247-248.]
26	Pan CD, Wang Q, Ruan X, Li ZH (2009) Biological activity and quantification of potential autotoxins from the leaves of Picea schtenkiana. Chinese Journal of Plant Ecology, 33, 186-196.(in Chinese with English abstract)
	[潘存德, 王强,阮晓, 李兆慧 (2009) 天山云杉针叶水提取物自毒效应及自毒物质的分离鉴定. 植物生态学报, 33, 186-196.]
27	Piao TF, Comita LS, Jin GZ, Kim JH (2013) Density dependence across multiple life stages in a temperate old-growth forest of Northeast China. Oecologia, 172, 207-217.
28	Qiu Y, Chang SL, Zhang YT, Wang WD, He P, Wang HJ, Xie J (2015) Biomass estimation modeling and adaptability analysis of organ allocation in six common shrub species in the forests of Tianshan Mountains. Acta Ecologica Sinica, 35, 7842-7851.(in Chinese with English abstract)
	[仇瑶, 常顺利, 张毓涛, 王文栋, 何平, 王慧杰, 谢锦 (2015) 天山林区六种灌木生物量的建模及其器官分配的适应性分析. 生态学报, 35, 7842-7851.]
29	Queenborough SA, Burslem D, Garwood NC, Valencia R (2007) Habitat niche partitioning by 16 species of Myristicaceae in Amazonian Ecuador. Plant Ecology, 192, 193-207.
30	Ripley BD (1977) Modelling spatial patterns. Journal of the Royal Statistical Society Series B: Statistical Methodology, 39, 172-212.
31	Stoll P, Newbery DM (2005) Evidence of species-specific neighborhood effects in the Dipterocarpaceae of a Bornean rain forest. Ecology, 86, 3048-3062.
32	Wang B, Huang YS, Li XK, Xiang WS, Ding T, Huang FZ, Lu SH, Han WH, Wen SJ, He LJ (2014) Species composition and spatial distribution of a 15 ha northern tropical karst seasonal rain forest dynamics study plot in Nonggang, Guangxi, southern China. Biodiversity Science, 22, 141-156.(in Chinese with English abstract)
	[王斌, 黄俞淞, 李先琨, 向悟生, 丁涛, 黄甫昭, 陆树华, 韩文衡, 文淑均, 何兰军 (2014) 弄岗北热带喀斯特季节性雨林15 ha监测样地的树种组成与空间分布. 生物多样性, 22, 141-156.]
33	Wang BS (1987) Phytocoenology. Higher Education Press, Beijing.(in Chinese)
	[王伯荪 (1987). 植物群落学. 高等教育出版社, 北京.]
34	Wang L, Sun QW, Hao CY, Tian SN, Zhang SS, Chen YK, Zhang XP (2010) Point pattern analysis of different age-class Taxus chinensis var. mairei individuals in mountainous area of southern Anhui Province. Chinese Journal of Applied Ecology, 21, 271-278.(in Chinese with English abstract)
	[王磊, 孙启武, 郝朝运, 田胜尼, 张姗姗, 陈一锟, 张小平 (2010) 皖南山区南方红豆杉种群不同龄级立木的点格局分析. 应用生态学报, 21, 271-278.]
35	Wang T, Ren SY, Yuan ZL, Zhu Y, Pan NL, Lu X, Ye YZ (2014) Effects of density dependence on the spatial patterns of Quercus aliena var. acuteserrata trees in deciduous broad-leaved forest in the Baotianman National Nature Reserve, Central China. Biodiversity Science, 22, 449-457.(in Chinese with English abstract)
	[王婷, 任思远, 袁志良, 祝燕, 潘娜李, 鹿鑫, 叶永忠 (2014) 密度制约对宝天曼落叶阔叶林锐齿栎死亡前后分布格局的影响. 生物多样性, 22, 449-457.]
36	Wang W, Rao MD, Chen SW, Zhu DH, Mi XC, Zhang JT (2014) Effects of negative density dependence and habitat filtering on temporal variation in phylogenetic community structure of seedlings in a mid-subtropical forest. Chinese Science Bulletin, 59, 1844-1850.(in Chinese with English abstract)
	[王薇, 饶米德, 陈声文, 朱大海, 米湘成, 张金屯 (2014) 负密度制约和生境过滤对古田山幼苗系统发育多样性时间变化的影响. 科学通报, 59, 1844-1850.]
37	Wang XG, Wiegand T, Wolf A, Howe R, Davies S, Hao ZQ (2011) Spatial patterns of tree species richness in two temperate forests. Journal of Ecology, 99, 1382-1393.
38	Wang XG, Hao ZQ, Ye J, Zhang J, Li BH, Yao XL (2008) Relationships 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, 27, 145-150.(in Chinese with English abstract)
	[王绪高, 郝占庆, 叶吉, 张健, 李步杭, 姚晓琳 (2008) 长白山阔叶红松林物种多度和空间分布格局. 生态学杂志, 27, 145-150.]
39	Wang XT, Hou YL, Liang CZ, Wang W, Liu F (2012) Point pattern analysis based on different null models for detecting spatial patterns. Biodiversity Science, 20, 151-158.(in Chinese with English abstract)
	[王鑫厅, 侯亚丽, 梁存柱, 王炜, 刘芳 (2012) 基于不同零模型的点格局分析. 生物多样性, 20, 151-158.]
40	Wang YH, Mi XC, Chen SW, Li MH, Yu MJ (2011) Regeneration dynamics of major tree species during 2002-2007 in a subtropical evergreen broad-leaved forest in Gutianshan National Nature Reserve in East China. Biodiversity Science, 19, 178-189.(in Chinese with English abstract)
	[汪殷华, 米湘成, 陈声文, 李铭红, 于明坚 (2011) 古田山常绿阔叶林主要树种2002-2007年间更新动态. 生物多样性, 19, 178-189.]
41	Wiegand T, Moloney KA (2004) Rings, circles and null- models for point pattern analysis in ecology. Oikos, 104, 209-229.
42	Zhang J, Hao ZQ, Sun IF, Song B, Ye J, Li BH, Wang XG (2009) Density dependence on tree survival in an old-growth temperate forest in northeastern China. Annals of Forest Science, 66(2), 1-9.
43	Zhang JT (1995) Methods in Vegetation Quantitative Ecology. China Science and Technology Press, Beijing.(in Chinese)
	[张金屯 (1995) 植被数量生态学方法. 中国科学技术出版社, 北京.]
44	Zhang L, Wang XJ, Hu EC, Gao GY, Pang X, Yu YZ, Zhang ZH (2011) Population structure and spatial pattern of Ulmus macrocarpa var. mongolica in Horqin Sandy Land, China. Journal of Desert Research, 31, 115-119.(in Chinese with English abstract)
	[张雷, 王晓江, 胡尔查, 高桂英, 庞鑫, 于有忠, 张振华 (2011) 科尔沁沙地蒙古黄榆种群结构与空间分布格局. 中国沙漠, 31, 115-119.]
45	Zhang M, Mi XC, Jin GZ (2014) Composition and spatial patterns of the Liangshui spruce-fir valley forest in the Xiao Hinggan Mountains. Chinese Science Bulletin, 59, 2377-2387.(in Chinese with English abstract)
	[张觅, 米湘成, 金光泽 (2014) 小兴安岭凉水谷地云冷杉林群落组成与空间格局. 科学通报, 59, 2377-2387.]
46	Zhang YT, Chang SL, Lu JJ, Li X, Wang Z, Shi QD, Zhang XP (2011a) Large scale permanent plot developed and its three-dimension realized in Tianshan forest. Scientia Silvae sinicae, 47, 179-183.(in Chinese with English abstract)
	[张毓涛, 常顺利, 芦建江, 李翔, 王智, 师庆东, 张新平 (2011a) 天山云杉森林8 ha样地的建立及三维可视化管理. 林业科学, 47, 179-183.]
47	Zhang YT, Li JM, Chang SL, Li X, Lu JJ (2011b) Spatial distribution pattern of Picea schrenkiana var. tianshanica population and its relationships with topographic factors in middle part of Tianshan Mountain.Chinese Journal of Applied Ecology, 22, 2799-2806.(in Chinese with English abstract)
1	Aminem E, Chang SL, Zhang YT, Qiu Y, He P (2014) Altitudinal distribution rule of Picea schrenkiana forest’s soil organic carbon and its influencing factors. Acta Ecologica Sinica, 34, 1626-1634.(in Chinese with English abstract)
	[阿米娜木·艾力, 常顺利, 张毓涛, 仇瑶, 何平(2014) 天山云杉森林土壤有机碳沿海拔的分布规律及其影响因素. 生态学报, 34, 1626-1634.]
47	[张毓涛, 李吉玫, 常顺利, 李翔, 芦建江 (2011b) 天山中部天山云杉种群空间分布格局及其与地形因子的关系. 应用生态学报, 22, 2799-2806.]
48	Zhang ZC, Hao ZQ, Ye J, Lin F, Yuan ZQ, Xing DL, Shi S, Wang XG (2013) Short-term death dynamics of trees in natural secondary poplar-birch forest in Changbai Mountains of Northeast China. Chinese Journal of Applied Ecology, 24, 303-310.(in Chinese with English abstract)
	[张昭臣, 郝占庆, 叶吉, 蔺菲, 原作强, 邢丁亮, 师帅, 王绪高 (2013) 长白山次生杨桦林树木短期死亡动态. 应用生态学报, 24, 303-310.]
49	Zhu Y, Mi XC, Ma KP (2009) A mechanism of plant species coexistence: the negative density-dependent hypothesis. Biodiversity Science, 17, 594-604.(in Chinese with English abstract)
2	Bagchi R, Henrys PA, Brown PE, Burslem DFRP, Diggle PJ, Savitri GCV, Nimal GIAU, Kassim AR, Law R, Noor S, Valencia RL (2011) Spatial patterns reveal negative density dependence and habitat associations in tropical trees. Ecology, 92, 1723-1729.
3	Chen XR, Chen YY, Luo ZR, Ding BY (2013) A 5-year mid-mountain subtropical evergreen broad-leaved forest study in Baishanzu, East China. Journal of Zhejiang A & F University, 30, 821-829.(in Chinese with English abstract)
49	[祝燕, 米湘成, 马克平 (2009) 植物群落物种共存机制: 负密度制约假说. 生物多样性, 17, 594-604.]
50	Zhu Y, Bai F, Liu HF, Li WC, Li L, Li GQ, Wang SZ, Sang WG (2011) Population distribution patterns and interspecific spatial associations in warm temperate secondary forests, Beijing. Biodiversity Science, 19, 252-259.(in Chinese with English abstract)
	[祝燕, 白帆, 刘海丰, 李文超, 李亮, 李广起, 王顺忠, 桑卫国 (2011) 北京暖温带次生林种群分布格局与种间空间关联性. 生物多样性, 19, 252-259.]
51	Zhu Y, Mi XC, Ren HB, Ma KP (2010) Density dependence is prevalent in a heterogeneous subtropical forest. Oikos, 119, 109-119.
3	[陈小荣, 陈圆圆, 骆争荣, 丁炳扬 (2013) 百山祖中山中亚热带常绿阔叶林群落5年动态特征. 浙江农林大学学报, 30, 821-829.]
4	Clark JS, Silman M, Kern R, Macklin E, Hille RJ (1999) Seed dispersal near and far: patterns across temperate and tropical forests. Ecology, 80, 1475-1494.
5	Connell JH (1971) On the role of natural enemies in preventing competitive exclusion in some marine animals and in rain forest trees. In: Dynamic of Populations (eds den Boer PJ, Gradwell G), pp. 298-312. Center Ag Publishing & Documentation Wageningen, Wageningen,.
6	Chen L, Mi XC, Comita L, Zhang LW, Ren HB, Ma KP (2010) Community-level consequences of density dependence and habitat association in a subtropical broad-leaved forest. Ecology Letters, 13, 695-704.
7	Condit R (1998) Ecological implications of changes in drought patterns: shifts in forest composition in Panama. Climatic Change, 39, 413-427
8	Condit R, Ashton PS, Baker P, Bunyavejchewin S, Gunatilleke S, Gunatilleke N, Hubbel SP, Foster RB, Itoh A, LaFrankie JV, Lee HS, Losos E, Manokaran N, Sukumar R, Yanakura T (2000) Spatial patterns in the distribution of tropical tree species. Science, 288, 1414-1418.
9	Condit R, Ashton PS, Manokaran N, LaFrankie JV, Hubbell SP, Foster RB (1999) Dynamics of the forest communities at Pasoh and Barro Colorado: comparing two 50-ha plots. Philosophical Transactions of the Royal Society of London B: Biological Sciences, 354, 1739-1748.
10	Coomes DA, Duncan RP, Allen RB, Truscott J (2003) Disturbances prevent stem size-density distributions in natural forests from following scaling relationships. Ecology Letters, 6, 980-989.
11	Diggle PJ, Chetwynd AG (1991) Second order analysis of spatial clustering for inhomogeneous populations. Biometrics, 47, 1155-1163.
12	Getzin S, Wiegand T, Wiegand K, He FL (2008) Heterogeneity influences spatial patterns and demographics in forest stands. Journal of Ecology, 96, 807-820.
13	Gunton RM, Kunin WE (2009) Density-dependence at multiple scales in experimental and natural plant populations. Journal of Ecology, 97, 567-580.

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