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

doi:10.3724/SP.J.1003.2014.14101

生物多样性 ›› 2014, Vol. 22 ›› Issue (4): 449-457. DOI: 10.3724/SP.J.1003.2014.14101

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

王婷¹, 任思远¹, 袁志良², 祝燕³, 潘娜¹, 李鹿鑫¹, 叶永忠^2,,A;^*()

1.河南农业大学林学院, 郑州 450002
2 .河南农业大学生命科学学院, 郑州 450002
3.中国科学院植物研究所植被与环境变化国家重点实验室, 北京 100093

收稿日期:2014-05-26 接受日期:2014-07-16 出版日期:2014-07-20 发布日期:2014-07-24
通讯作者: 叶永忠
基金资助:
国家自然科学基金(31270493)和植被与环境变化国家重点实验室2012年开放课题(LVEC-2012kf06)

Effects of density dependence on the spatial patterns of Quercus aliena var. acuteserrata trees in deciduous broad-leaved forest in the Baotianman Nature Reserve, central China

Ting Wang¹, Siyuan Ren¹, Zhiliang Yuan², Yan Zhu³, Na Pan¹, Luxin Li¹, Yongzhong Ye^2,^*()

1. Forestry College, Henan Agricultural University, Zhengzhou 450002
2. College of Life Sciences, Henan Agricultural University, Zhengzhou 450002
3. State Key Laboratory of Vegetation and Environmental Change, Institute of Botany, Chinese Academy of Science, Beijing 100093

Received:2014-05-26 Accepted:2014-07-16 Online:2014-07-20 Published:2014-07-24
Contact: Ye Yongzhong

摘要/Abstract

摘要：

为了在温带和亚热带的过渡带中验证森林树木死亡是否受密度制约的影响, 我们在宝天曼国家级自然保护区选择1个100 m×100 m的暖温带落叶阔叶林样地, 用双变量函数g(r)(the pair-correlation function)研究了锐齿栎(Quercus aliena var. acuteserrata)死亡前(活树和枯木统称为死亡前树木)和死亡后(活树为死亡后树木)的分布格局。把样地中的锐齿栎分为幼树(1 cm ≤ DBH<10 cm)、小树(10 cm ≤ DBH<20 cm)和成年树(DBH≥20 cm)3个不同的生长阶段来研究密度制约对空间分布格局的影响机制。结果表明: (1)死亡前锐齿栎在r>5 m尺度呈聚集分布, 死亡后幼树(1 cm≤DBH<10 cm)和成年树(DBH≥20 cm)在1-25 m尺度呈现随机分布, 死亡后小树(10 cm≤DBH< 20 cm)在r<1.5 m和2.5-4.5 m的尺度为随机分布, 在r>5 m的尺度呈聚集分布; (2)采用随机标签零模型和案例-对照设计的方法, 排除生境异质性影响后, 将幼树和小树的分布格局作为案例, 将成年树的分布格局作为对照, 并代表生境异质性的作用, 通过小径级树木与成年树分布格局的对比发现, 密度制约效应对死亡前后的锐齿栎分布格局均具有影响; (3)幼树和小树在成年树周围的分布死亡前为显著聚集分布格局, 死亡后剩余树木的聚集强度下降; 随着与成年树之间距离的增加, 死亡后的幼树(仅包括现存的活树)逐渐向随机分布格局演替。本研究初步表明锐齿栎空间格局受生境异质性的影响并呈现出显著的聚集效应, 排除生境异质性影响后, 锐齿栎死亡前后的空间格局受到密度制约的影响, 这一结果为Janzen-Connell假说提供了支持。

关键词: 落叶阔叶林, Quercus aliena var. acuteserrata, 密度制约, 分布格局, Janzen-Connell假说, 宝天曼国家级自然保护区

Abstract

To determine the contribution of density dependence to tree mortality in the transitional region between temperate and subtropical zone, a deciduous broad-leaf forest plot (100 m×100 m) in the Baotianman National Nature Reserve was selected and a pair-correlation function g(r) (the pair-correlation function) was employed to examine the spatial pattern of a single species. Individuals of the dominant species Quercus aliena var. acuteserrata were divided into three growth stages: saplings (1 cm ≤ DBH < 10 cm), juveniles (10 cm ≤ DBH < 20 cm), and adult trees (DBH ≥ 20 cm). Each stage was then divided into pre-mortality (including all living and dead trees) and post-mortality (only living trees) status to examine the contribution of density dependence to the spatial patterns of Q. aliena var. acuteserrata. The results showed that: (1) Pre-mortality Q. aliena var. acuteserrata trees showed an aggregated distribution pattern at r > 5 m scale. Post-mortality Q. aliena var. acuteserrata saplings and adult trees had a random distribution pattern at a 1-25 cm scale. Post-mortality juveniles had a random distribution pattern at r < 1.5 m and 2.5-4.5 m scales and an aggregated distribution pattern at r > 5 m scale. (2) The spatial pattern of adult trees was regarded as a control pattern accounting for environmental heterogeneity. The spatial pattern of pre- and post-mortality saplings and juveniles showed density-dependent distribution responses by random labeling null model with a case-control design; (3) Pre-mortality saplings and juveniles showed a clumped distribution around adult trees, whereas the post-mortality saplings and juveniles displayed a weak clustering with mortality caused by density dependence. With increasing distance to adult trees, post-mortality saplings and juveniles showed a gradually increasing random distribution. Our findings indicate that habitat heterogeneity contributes to the spatial distribution of Q. aliena var. acuteserrata with an aggregation effect. After the effect of habitat heterogeneity was removed, spatial distribution of Q. aliena var. acuteserrata trees with different pre-mortality and post-mortality status were all affected by density dependence effect in the Baotianman National Nature Reserve. These conclusions provide support for the Janzen-Connell hypotheses.

Key words: deciduous broad-leaf forest, Quercus aliena var. acuteserrata, density dependence, distribution pattern, Janzen-Connell hypotheses, Baotianman National Nature Reserve

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. DOI: 10.3724/SP.J.1003.2014.14101.

Ting Wang, Siyuan Ren, Zhiliang Yuan, Yan Zhu, Na Pan, Luxin Li, Yongzhong Ye (2014) Effects of density dependence on the spatial patterns of Quercus aliena var. acuteserrata trees in deciduous broad-leaved forest in the Baotianman Nature Reserve, central China. Biodiversity Science, 22, 449-457. DOI: 10.3724/SP.J.1003.2014.14101.

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链接本文: https://www.biodiversity-science.net/CN/10.3724/SP.J.1003.2014.14101

https://www.biodiversity-science.net/CN/Y2014/V22/I4/449

图/表 5

图1 宝天曼1 ha样地(100 m×100 m)中锐齿栎枯木的点格局分析。黑色实线为实际点格局, 虚线之间为置信区间。

Fig. 1 Spatial patterns of Quercus aliena var. acuteserrata deadwoods in the 1-ha plot in the Baotianman National Nature Reserve. The solid black line represents the point pattern of the dead trees, the confidence interval is showed by the region between the two dashed gray lines.

图2 宝天曼1 ha样地锐齿栎活树的分布格局, 基于完全随机模型下的双关联函数g(r)和L(r)函数的格局分析。黑色实线表实际点的格局, 灰色区间和虚线之间代表置信区间。

Fig. 2 Spatial patterns of living Q. aliena var. acuteserrata trees in the 1-ha plot in Baotianman Nature Reserve. It presents the double correlation functions g(r) and L(r) functions pattern analysis with complete spatial randomness model. The solid black line represents the point pattern of the living trees, the confidence interval is shown by the gray part and the region between the two dashed lines.

图3 宝天曼1 ha样地不同径级间锐齿栎枯木点格局分析, A为幼树, B为小树, C为成年树。黑实线表实际点的格局, 虚线之间代表置信区间。

Fig. 3 Point pattern analysis of Q. aliena var. acuteserrata deadwoods at different growth stages in the 1-ha plot in the Baotianman National Nature Reserve. The solid black line represents the point pattern of the dead trees, the confidence interval is shown by the region between the dotted lines. A, Saplings (1 cm ≤ DBH < 10 cm); B, Juveniles (10 cm ≤ DBH <20 cm); C, Adult trees (DBH ≥20 cm).

图4 宝天曼锐齿栎密度制约效应分析。以成年树的格局作为对照代表生境异质性, 幼树或小径级树木的格局作为案例格局, a为幼树, b为小树, c为成年树; A1、B1分别为死亡前幼树、小树为案例的密度制约效应分析, A2、B2分别为死亡后幼树、小树为案例的密度制约效应分析。

Fig. 4 Examples for density dependent effect within a case-control design in the 1-ha plot in the Baotianman National Nature Reserve. The pattern of adult trees serves as “control”, which corrects for possible heterogeneity in habitat quality, the pattern of smaller size classes serves as “cases”. A, Saplings (1 cm ≤ DBH <10 cm); B, Juveniles (10 cm ≤ DBH < 20 cm); C, Adult trees (DBH ≥ 20 cm). A1 and B1 represent the negative density dependence analysis of pre-mortality saplings and juveniles; A2 and B2 show the negative density dependence analysis of post-mortality saplings and juveniles, respectively.

图5 运用双关联函数g12(r)和独立性零模型分析不同生长阶段锐齿栎死亡前后的关联性. A1到F1为死亡前, A2到F2为死亡后。a为幼树, b为小树, c为成年树。Monte Carlo模拟循环999次取99%的置信区间(灰色区间)。

Fig. 5 Comparative analysis of the correlation of pre-mortality and post-mortality of Quercus aliena var. acuteserrata trees at different growth stages by using double correlation functions g12 (r) and random labeling null model. A1to F1 for pre-mortality trees, A2 to F2 for post-mortality trees. a, Saplings (1 cm ≤ DBH <10 cm); b, Juveniles (10 cm ≤ DBH <20 cm); c, Adult trees (DBH ≥20 cm). It shows the 99% confidence interval (grey section) after 999 times circles with Monte Carlo simulation.

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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

Effects of density dependence on the spatial patterns of Quercus aliena var. acuteserrata trees in deciduous broad-leaved forest in the Baotianman Nature Reserve, central China

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