植物生态学报, 2014, 38(6): 576-584 doi: 10.3724/SP.J.1258.2014.00053

研究论文

海南岛霸王岭国家重点保护植物的生态位研究

陈玉凯1,*, 杨琦1,*, 莫燕妮2, 杨小波,1,**, 李东海1, 洪小江3

1海南大学热带作物种质资源保护与开发利用教育部重点实验室, 海口 570228

2海南省林业厅, 海口 570203

3海南霸王岭国家级自然保护区, 海南昌江 572722

A study on the niches of the state’s key protected plants in Bawangling, Hainan Island

CHEN Yu-Kai1,*, YANG Qi1,*, MO Yan-Ni2, YANG Xiao-Bo,1,**, LI Dong-Hai1, HONG Xiao-Jiang3

1Key Laboratory of Protection and Development Utilization of Tropical Crop Germplasm Resources, Ministry of Education, Hainan University, Haikou 570228, China

2Forestry Department of Hainan Province, Haikou 570203, China

3Bawangling National Nature Reserve, Changjiang, Hainan 572722, China

通讯作者: ** (E-mail: yanfengxb@163.com)

第一联系人:

* 同等贡献, 并列第一作者。These authors contributed equally to this work and should be considered co-first authors.

编委: 张金屯

责任编辑: 王 葳

收稿日期: 2014-01-27   接受日期: 2014-04-10   网络出版日期: 2014-06-01

基金资助: 国家自然科学基金.  31060073
海南省第二次全国重点保护植物资源调查项目

Corresponding authors: * (E-mail: yanfengxb@163.com)

Received: 2014-01-27   Accepted: 2014-04-10   Online: 2014-06-01

摘要

为进一步了解国家重点野生保护植物的生存群落及不同种群利用资源和占据生态空间的能力, 为制定国家重点保护植物种群的保护措施提供依据, 2009-2011年, 对霸王岭有国家重点保护植物分布的70个样地进行群落调查, 研究了霸王岭保护区内12种国家重点保护植物的生态位宽度以及物种间的生态位重叠。结果表明: (1)生态位宽度值从大到小依次为: 土沉香(Aquilaria sinensis)、海南韶子(Nephelium topengii)、普洱茶(Camellia sinensis var. assamica)、油丹(Alseodaphne hainanensis)、海南油杉(Keteleeria hainanensis)、鸡毛松(Podocarpus imbricatus)、海南梧桐(Firmiana hainanensis)、乐东拟单性木兰(Parakmeria lotungensis)、海南粗榧(Cephalotaxus mannii)、香子含笑(Michelia hedyosperma)、粘木(Ixonanthes reticulata)和陆均松(Dacrydium pierrei); (2)物种生态位宽度值的大小与其自身的地理分布范围密切相关, 如: 陆均松、乐东拟单性木兰等生态位宽度较小的原因与其分布的海拔范围狭窄有关。生态位宽度较大的种群对资源的利用能力较强, 与其他物种的生态位重叠一般较大, 如: 生态位宽度较大的普洱茶与海南油杉的生态位重叠值最大, 为0.872, 而一些生态位宽度较小的物种生态位重叠值较高, 如: 香子含笑的生态位宽度较小, 但与海南梧桐的生态位重叠较大, 为0.693。因此, 种群间的生态位重叠值与它们对环境资源利用的相似程度以及物种自身的生物学特性密切相关; (3)香子含笑、粘木等的生态位宽度较小, 而且个体数量较少, 适宜其生存的群落和生境极少, 应该给予更多的关注和优先保护; (4)未来对濒危植物的保护不仅要考虑各物种自身的生物学特性, 还应从植物群落的角度考虑。只有保护好这些濒危植物的生存群落, 如山地雨林等, 才能实现对濒危植物的有效保护。

关键词: 霸王岭 ; 群系 ; 国家重点保护植物 ; 生态位

Abstract

Aims An understanding on the living communities of the state’s key protected wild plants and the ability in utilizing resources and occupying ecological spaces by different populations is essential to the protection and propagation of rare and endangered plant species. We use our findings to highlight the knowledge gaps in the conservation strategies for the endangered species.
Methods Field investigations were conducted during 2009-2011. Based on the survey data on 70 forest plots in the Bawangling protected area, the niche breadths and overlaps of 12 species listed as the state’s key protected plants were calculated and analyzed.
Important findings The 12 species, by their niche breadth values in descending order, include Aquilaria sinensis, Nephelium topengii, Camellia sinensis var. assamica, Alseodaphne hainanensis, Keteleeria hainanensis, Podocarpus imbricatus, Firmiana hainanensis, Parakmeria lotungensis, Cephalotaxus mannii, Michelia hedyosperma, Ixonanthes reticulata, Dacrydium pierrei. The niche breadth value of a species is closely related to its range of geographical distribution; both Dacrydium pierrei and Parakmeria lotungensis had a narrow niche breadth corresponding to their narrow ranges of altitudinal distribution. In general, the niche overlap value is higher between species with greater niche breadth; the highest niche overlap value of 0.872 was found between Camellia sinensis var. assamica and Keteleeria hainanensis. However, the niche overlap value between species with smaller niche breadth could also be high; e.g. the niche overlap value was relatively high (0.693) between two species with smaller niche breadth, Michelia hedyosperma and Firmiana hainanensis, indicating that niche overlap value is dependent upon the biological characteristics and requirements for environmental resources of the species of comparison. Some species with low niche breadth values (e.g. Michelia hedyosperma and Ixonanthes reticulata) were found to have smaller populations. Due to lack of suitable community and habitats, these plant species are in dire need of more attention and prioritization for conservation. In conclusion, for the better protection and propagation of these endangered plants in the future, we should combine knowledge of their biological characteristics with that of their living communities such as the montane rainforests, etc.

Keywords: Bawangling ; formation ; national protected plant ; niche

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本文引用格式

陈玉凯, 杨琦, 莫燕妮, 杨小波, 李东海, 洪小江. 海南岛霸王岭国家重点保护植物的生态位研究. 植物生态学报[J], 2014, 38(6): 576-584 doi:10.3724/SP.J.1258.2014.00053

CHEN Yu-Kai, YANG Qi, MO Yan-Ni, YANG Xiao-Bo, LI Dong-Hai, HONG Xiao-Jiang. A study on the niches of the state’s key protected plants in Bawangling, Hainan Island. Chinese Journal of Plant Ecology[J], 2014, 38(6): 576-584 doi:10.3724/SP.J.1258.2014.00053

生态位(niche)可以定量地研究物种之间的关系、物种和环境之间的相互关系, 已成为现代生态学研究的一个重要课题, 特别是在种间关系、群落结构、群落演替、生物多样性物种进化等方面的研究中获得了广泛的应用(Schoener, 1970; 王刚等, 1984; Mueller & Altenberg, 1985; Westman, 1991; 刘金福和洪伟, 1999; 李登武等, 2005; 王正宁等, 2005), 成为解释自然群落中物种共存与竞争机制的基本理论。因此, 生态位在研究生物多样性的保护及保护物种的评价方面具有较高的生态学价值, 反映了种群对资源和生态空间的利用能力, 同时也是不同的种群适应不同环境的生存策略和机制(Robert et al., 2003)。

海南霸王岭国家级自然保护区位于海南岛西南部山区, 保护区的主要植被类型是热带雨林, 而热带雨林作为生态系统重要的组成部分, 拥有最高的物种多样性。热带雨林的生物多样性导致了热带雨林群落结构的复杂性, 因此, 对于热带地区的重点野生保护植物的调查与研究显得尤为重要。目前国内外对该地区热带雨林群落的研究主要集中在植物多样性和优势种的群落结构特征等方面(胡玉佳和丁小球, 2000; 臧润国等, 2001, 2003; 张志东和臧润国, 2007; 刘万德等, 2009)。目前, 有关该区国家重点野生保护植物在整个保护区不同群落中利用资源和占据生态空间能力的研究尚未见报道。

由于人为对环境的破坏、生态旅游的开发等, 许多濒危植物所处的生境面临着较大的威胁, 虽然近年来对濒危植物的生存状态、种群生态位特征, 以及它们对空间资源的利用及其濒危机制等的研究逐渐增多(史红文等, 2007; 史小华等, 2007; 王立龙等, 2007; 刘春生等, 2009), 但这些研究主要围绕一种植物的生存状态和保护对策进行综合分析, 从生态位角度探讨不同濒危植物的种间竞争及共存特征、濒危机制等, 进而提出一种普适性的对策和方法的研究较少。那么, 在同一环境中不同濒危植物是如何相互适应的?它们对资源的利用途径以及在不同环境下的生态适应性是否存在着一些共同特征?这两个科学问题有待较多的研究案例来论证。为此, 本文通过对霸王岭保护区国家重点保护植物在不同群落中的生态位状况进行分析, 探讨不同国家重点保护植物种群之间的相互关系, 为制定国家重点保护植物种群的保护措施提供理论依据。

1 材料和方法

1.1 研究地区概况

海南霸王岭国家级自然保护区位于海南岛西南部山区, 地理坐标为18.87°-19.20° N, 108.88°- 109.33° E, 属于森林生态系统类型的自然保护区。保护区总面积29β980 hm2, 核心区面积为10β540 hm2, 缓冲区面积为8β910 hm2, 实验区面积为10β530 hm2。该地区属于热带季风气候, 干湿季节较明显。海拔高度在100-1700 m范围, 保护区内的土壤以沙或花岗岩作为母质而发育成为砖红壤为代表的类型, 随着海拔高度的增加不断过渡为山地红壤(胡玉佳和丁小球, 2000)。低地雨林和山地雨林是该地区分布最广的两个主要植被类型, 低地雨林是以青梅(Vatica mangachapoi)、荔枝(Litchi chinensis)等为优势种的植被类型, 人为干扰强度较大。以鸡毛松(Podocarpus imbricatus)、陆均松(Dacrydium pierrei)等为优势种的山地雨林分布海拔较高, 人为干扰强度较小, 因而至今仍保存一定面积的原始山地雨林植被。霸王岭林区大部分的原始森林都经过了刀耕火种和商业性采伐的破坏, 自1994年海南岛实施全岛森林禁伐以来, 霸王岭的热带森林大部分处于自然恢复状态(张志东和臧润国, 2007)。

1.2 研究方法

1.2.1 样地调查

在霸王岭保护区内沿着不同的海拔、坡向、坡度和不同管护点或监测点设置样地, 根据2011年国家林业局对中国重点保护野生植物资源调查的技术方法(国家林业局, 2009), 对乔木树种及大灌木的样方(面积为400 m2, 即每个样地为20 m × 20 m)同时记录各样地的海拔、坡度、坡向等生境指标, 将每个样地再划分成5 m × 5 m的小样方, 对其进行每木胸径、枝下高、树高、冠幅等调查。共设置70个样地, 总面积2.8 hm2。同时, 对分布在大面积森林群落中且出现于3个以上不同小生境中的海南梧桐(Firmiana hainanensis)、香子含笑(Michelia hedyosperma)、陆均松、粘木(Ixonanthes reticulata)、鸡毛松、海南粗榧(Cephalotaxus mannii)、海南油杉(Keteleeria hainanensis)、油丹(Alseodaphne hainanensis)、乐东拟单性木兰(Parakmeria lotungensis)、海南韶子(Nephelium topengii)、土沉香(Aquilaria sinensis)、普洱茶(Camellia sinensis var. assamica)等12种国家重点保护植物的生态位进行研究, 12个物种所分布的群落可划分为14个不同的群系, 具体为: 厚皮香八角(Illicium ternstroemioides) +厚壳桂(Cryptocarya chinensis)群系、陆均松+乐东拟单性木兰群系、海南白椎(Castanopsis carlesii var. hainanica) +九节(Psychotria asiatica)群系、红柯(Lithocarpus fenzelianus) +谷木(Memecylon ligustrifolium)群系、厚壳桂+药用狗牙花(Tabernaemontana bovina)群系、油丹+黄叶树(Xanthophyllum hainanensis)群系、线枝蒲桃(Syzygium araiocladum) +九节群系、厚壳桂+线枝蒲桃群系、海南山龙眼(Helicia hainanensis) +厚壳桂群系、海南倒吊笔(Wrightia laevis) +百两金(Ardisia crispa)群系、药用狗牙花+黄丹木姜子(Litsea elongata)群系、白背厚壳桂(Cryptocarya maclurei) +药用狗牙花群系、海南韶子+鹅掌柴(Schefflera octophylla)群系、重阳木(Bischofia javanica) +阴香(Cinnamomum burmanni)群系。本文以这14个群系作为“资源状态”, 计算这12种国家重点保护植物在不同群系中的重要值, 进而得出它们的生态位宽度以及生态位重叠度, 并分析它们在不同群落中的地位及相互之间利用资源的状况。

1.2.2 数据分析方法

1.2.2.1 物种重要值的计算 采用《植物群落实验手册》上的常规分析方法统计各样地出现的物种数、个体数、频度、物种出现的样方数, 计算每个种群的重要值,

IV(重要值) = (相对多度+相对频度+相对显著度)/3 (王伯荪等, 1996)。

其中, 相对多度指某个种在全部样方中的个体数与所有种个体总和之比, 即相对多度=某个种的个体数/所有种的个体数×100%; 相对频度指某个种在全部样方中的频度与所有种频度总和之比, 即相对频度=某个种的频度值/所有种的频度值×100%; 相对显著度指某个种在全部样方中的胸高断面积与所有种胸高断面积总和之比, 即相对显著度= (某个种的胸高断面积/所有种的总胸高断面积) × 100%。

1.2.2.2 生态位的研究方法 选取重要值作为资源利用的参数, 由此得出的生态位宽度就具有多维的意义(王刚等, 1984)。本文以不同的保护植物所在的14个群系为“资源状态”, 重要值为计测的状态指标, 研究霸王岭主要保护物种的生态位。

(1)生态位宽度采用Shannon-Wiener计算公式(张金屯, 2004):

$B_{i}=-\sum_{j=1}^{r}(P_{ij} \ln P_{ij})$

式中, Bi为种i的生态位宽度; Pij= nij/Ni, 式中nij为物种i在资源j上的重要值, Ni表示物种i在所有资源上的重要值之和。Pij代表种i在资源j上的重要值占该种在所有资源上的重要值的比例, r为资源位总位数。

(2)生态位重叠

生态位重叠多认为是种群对相同资源的共同利用, 或者是共有的生态空间资源区域。本文选取了2种计算方式研究种群之间的生态位重叠:

a)生态位相似性比例系指两个物种利用资源的相似程度, 其计算公式(李登武等, 2005; 彭逸生等, 2007)为:

$C_{ih}=1-\frac{1}{2}\sum\limits_{j=1}^{r}|P_{ij}-P_{hj}|$

式中, Cih为物种i与物种h的相似性程度, 数值范围为[0,1]; PijPhj分别为种i和种h在资源j上的重要值百分率, 且Cih = Chi

b)生态位重叠系指一定资源序列上, 两个物种利用同等级资源且相互重叠的情况, 其计算公式(Pianka, 1973)为:

$O_{ik}=\sum\limits_{j=1}^{r}P_{ij}P_{kj}/\sqrt{\sum\limits_{j=1}^{r}P_{ij}^{2}\sum\limits_{j=1}^{r}P_{kj}^{2}}$

式中, PijPkj分别为种i和种k在资源为j上的重要值。

2 结果和分析

2.1 霸王岭国家重点保护植物重要值特征

表1可知, 这12种保护植物主要分布在线枝蒲桃+九节群系和白背厚壳桂+药用狗牙花群系等群落中。12种不同保护植物在不同群落中的重要值差异较显著, 如油丹, 主要分布在线枝蒲桃+九节群系、油丹+黄叶树群系、海南山龙眼+厚壳桂群系中, 而该种群在其他群落中的优势地位不明显; 鸡毛松和海南韶子在海南韶子+鹅掌柴群系中优势度明显, 海南油杉在厚壳桂+药用狗牙花群系中优势度明显; 一些物种不仅在以自身为优势种的群落中具有明显优势, 而且在其他群落中具有一定的优势, 如乐东拟单性木兰在陆均松+乐东拟单性木兰群系中优势度明显, 在海南白椎+九节群系和红柯+谷木群系中同样具有极大的优势。

表1   12种保护植物在不同群落中的重要值

Table 1  Importance value of the 12 protected plants in different plant communities

序号 No.ABCDEFGHIJKLMN
鸡毛松 Podocarpus imbricatus0.0000.0000.2080.5460.0000.0000.3830.3620.0000.0000.0001.3642.0960.000
海南油杉 Keteleeria hainanensis6.7050.0002.4761.2958.6480.5183.3520.0000.0000.0000.0000.0000.0000.000
油丹 Alseodaphne hainanensis0.0001.6920.0001.2050.43710.2276.78911.8989.3110.0000.1990.0000.9120.000
海南韶子 Nephelium topengii0.0000.0000.0000.2950.0003.5362.0021.3802.4580.6761.5390.5725.1090.000
土沉香 Aquilaria sinensis0.0000.1600.9890.0000.0000.7450.9210.1770.0000.6760.1980.8480.3580.000
普洱茶 Camellia sinensis
var. assamica
3.8490.2891.5260.0003.8071.4180.0001.1770.0000.0000.3981.5020.5550.000
海南梧桐 Firmiana hainanensis0.0000.0000.0000.0000.0000.0000.0000.0000.0005.1840.0003.6273.07410.927
乐东拟单性木兰
Parakmeria lotungensis
0.0006.26210.3792.3440.0000.1130.3270.0000.0000.0000.0000.0000.0000.000
香子含笑 Michelia hedyosperma0.0000.0000.0000.0000.0000.0000.0000.0000.0000.0000.2861.1810.0000.811
陆均松 Dacrydium pierrei0.00013.5850.0000.0000.0000.0004.0561.9080.0000.0000.0000.0000.0000.000
粘木 Ixonanthes reticulate0.0000.0000.2142.0700.0000.0001.8090.0000.0000.0000.0000.0000.0000.000
海南粗榧 Cephalotaxus mannii0.0000.0000.0000.0000.0000.0000.0000.0000.0006.56815.05610.5920.0000.000

A, 厚皮香八角+厚壳桂群系; B, 陆均松+乐东拟单性木兰群系; C, 海南白椎+九节群系; D, 红柯+谷木群系; E, 厚壳桂+药用狗牙花群系; F, 油丹+黄叶树群系; G, 线枝蒲桃+九节群系; H, 厚壳桂+线枝蒲桃群系; I, 海南山龙眼+厚壳桂群系; J, 海南倒吊笔+百两金群系; K, 药用狗牙花+黄丹木姜群系; L, 白背厚壳桂+药用狗牙花群系; M, 海南韶子+鹅掌柴群系; N, 重阳木+阴香群系。

A, community of Illicium ternstroemioides and Cryptocarya chinensis; B, community of Dacrydium pierrei and Parakmeria lotungensis; C, community of Castanopsis carlesii var. hainanica and Psychotria asiatica; D, community of Lithocarpus fenzelianus and Memecylon ligustrifolium; E, community of Cryptocarya chinensis and Tabernaemontana bovina; F, community of Alseodaphne hainanensis and Xanthophyllum hainanensis; G, community of Syzygium araiocladum and Psychotria asiatica; H, community of Cryptocarya chinensis and Syzygium araiocladum; I, community of Helicia hainanensis and Cryptocarya chinensis; J, community of Wrightia laevis and Ardisia crispa; K, community of Tabernaemontana bovina and Litsea elongata; L, community of Cryptocarya maclurei and Tabernaemontana bovina; M, community of Nephelium topengii and Schefflera octophylla; N, community of Bischofia javanica and Cinnamomum burmanni.

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2.2 生态位宽度

表2可知, 土沉香、海南韶子和普洱茶生态位宽度值较大, 分别为2.017、1.923和1.907, 表明这3个物种能够出现在霸王岭的多个群系中, 其生态适应力也相对其他保护植物强, 且在我们的实际调查中, 以上分布范围较广的保护物种, 在各种群系中都有植株存在, 反映出它们对不同环境的生态适应力。香子含笑、粘木和陆均松的生态位宽度值较小, 分别为0.969、0.859和0.806, 说明这3个物种在霸王岭地区分布的范围较窄, 且实际调查也表明这3个物种在空间分布上极不均匀, 反映出它们对群落生境的要求更为苛刻, 导致其适宜生存的生境也较少。同时我们发现, 生态位宽度较小的这3种植物在不同群落中的分布也较少, 仅出现在3个群系中, 如香子含笑分布的森林植被类型为药用狗牙花+黄丹木姜群系、白背厚壳桂+药用狗牙花群系和重阳木+阴香群系, 并且在这3种群落中分布的数量极少; 粘木分布的森林群落类型为海南白椎+九节群系、红柯+谷木群系和线枝蒲桃+九节群系, 分布的数量相对较多, 尤其以红柯+谷木群系分布的数量最多; 而陆均松相对于香子含笑和粘木分布的植株数量较多, 但在不同的林分中分布不均匀, 在厚皮香八角+厚壳桂群系的分布数量最多, 表明陆均松在厚皮香八角+厚壳桂群系的群落类型较适宜的生境中显示出种群优势, 但由于人为干扰及物种自身生物学和生态学特性, 使得其分布的地理范围变得越来越窄。

表2   12种保护植物的生态位宽度值

Table 2  Niche breadth of the 12 protected plants

序号
No.
物种
Species
Shannon-Wiener
生态位宽度
Shannon-Wiener niche breadth
序号
No.
物种
Species
Shannon-Wiener
生态位宽度
Shannon-Wiener niche breadth
1鸡毛松 Podocarpus imbricatus1.4847海南梧桐 Firmiana hainanensis1.252
2海南油杉 Keteleeria hainanensis1.4958乐东拟单性木兰 Parakmeria lotungensis1.054
3油丹 Alseodaphne hainanensis1.7069香子含笑 Michelia hedyosperma0.969
4海南韶子 Nephelium topengii1.92310陆均松 Dacrydium pierrei0.806
5土沉香 Aquilaria sinensis2.01711粘木 Ixonanthes reticulata0.859
6普洱茶 Camellia sinensis var. assamica1.90712海南粗榧 Cephalotaxus mannii1.045

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不同的物种在不适宜自身的不利环境条件下会选择不同的适应对策, 对不利环境条件的适应能力较强的物种, 可通过扩大它们的地理分布以保持有足够的植株数量, 而适应能力较差的物种只能选择在较适宜自身发展的环境中保持优势的个体数量来维持其种群的生存。

2.3 生态位相似性

表3可知, 12个物种间的生态位相似性比例值差异显著, 其中生态位相似性比例值在0.6以上的物种共11对, 占总数的16.7%, 其中生态位相似性比例值在0.8以上的有4对, 分别是鸡毛松和香子含笑、海南油杉和油丹、海南油杉和海南粗榧以及土沉香与香子含笑, 说明这几个物种之间对资源的利用程度基本一致; 而多数种对(31对, 占总数的46.9%)物种之间的生态位相似性比例值较小, 介于0.1-0.4之间, 说明多数物种对环境以及资源的要求差异性较大; 另外, 还有2个种对之间的生态位相似性比例值不到0.01, 如鸡毛松与海南油杉、土沉香与海南粗榧等, 说明这几个物种之间对环境的要求基本不同。同时, 生态位宽度较大的物种, 它们之间的生态位相似性比例值也较大, 如: 海南油杉与油丹、海南油杉与海南粗榧, 其生态位相似性比例值均在0.85以上; 而一些生态位宽度较小的物种之间的生态位相似性比例值有时要高于那些生态位宽度值较大的种对, 如香子含笑与鸡毛松之间的生态位相似性比例为0.804, 这主要与它们对环境资源利用的相似程度以及物种自身的生物学特性有关。

表3   12种保护植物的生态位相似性

Table 3  Niche similarity of the 12 protected plants

物种
Species
鸡毛松
Podocarpus imbricatus
海南油杉
Keteleeria hainanensis
油丹
Alseodaphne
hainanensis
海南韶子
Nephelium topengii
土沉香
Aquilaria sinensis
普洱茶
Camellia sinensis
var. assamica
海南梧桐
Firmiana
hainanensis
乐东拟单性木兰
Parakmeria
lotungensis
香子含笑
Michelia
hedyosperma
陆均松
Dacrydium
pierrei
粘木
Ixonanthes
reticulata
海南油杉 Keteleeria hainanensis0.004
油丹 Alseodaphne hainanensis0.4030.930
海南韶子 Nephelium topengii0.4810.3620.292
土沉香 Aquilaria sinensis0.7780.0710.4090.453
普洱茶 Camellia sinensis var. assamica0.4310.3370.6810.1010.520
海南梧桐 Firmiana hainanensis0.1770.7980.2130.1650.0880.295
乐东拟单性木兰 Parakmeria lotungensis0.2830.2030.7840.1890.3060.0520.516
香子含笑 Michelia hedyosperma0.8040.0110.4670.3560.8280.4690.1740.294
陆均松 Dacrydium pierrei0.3890.2140.4780.0980.4420.0790.3510.1990.459
粘木 Ixonanthes reticulate0.7330.1700.3450.3270.7200.2920.0690.4770.7410.437
海南粗榧 Cephalotaxus mannii0.0590.8620.3400.3950.0060.3820.2480.5790.0580.4140.132

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2.4 生态位重叠

表4可知, 虽然不同保护植物之间的生态位重叠值存在一定的差异, 但多数保护植物之间的生态位重叠值较小, 可见多数国家保护物种适宜的生境较少, 而且在适宜生态位资源方面表现出分异或趋同的特性。生态位宽度和生态位重叠之间存在较大的相关性。如, 由于海南韶子的生态位宽度较大, 与鸡毛松和油丹等物种的生态位重叠值也较大, 分别为0.672和0.656; 生态位宽度较大的普洱茶与海南油杉的生态位重叠值最大, 为0.872; 生态位宽度较小的香子含笑、陆均松、粘木、海南梧桐、乐东拟单性木兰以及海南粗榧与一些保护植物之间的生态位重叠值为0; 但有些生态位宽度较小的物种之间的生态位重叠却较大, 如: 香子含笑的生态位宽度较小, 但与海南梧桐和海南粗榧的生态位重叠分别为0.693和0.589, 而有些生态位宽度较大的物种的生态位重叠值却不高, 如海南油杉与鸡毛松的生态位重叠为0.081, 说明生态位宽度较大的物种虽然对资源的利用能力较强, 分布较广, 但与其他种群间的生态位重叠不一定大。生态位宽度较小的物种, 与其他种群间的生态位重叠值也不一定小。

表4   12种保护植物的生态位重叠

Table 4  Niche overlap of the 12 protected plants

物种
Species
鸡毛松
Podocarpus imbricatus
海南油杉
Keteleeria hainanensis
油丹
Alseodaphne
hainanensis
海南韶子
Nephelium topengii
土沉香
Aquilaria sinensis
普洱茶
Camellia sinensis
var. assamica
海南梧桐
Firmiana
hainanensis
乐东拟单性木兰
Parakmeria
lotungensis
香子含笑
Michelia
hedyosperma
陆均松
Dacrydium
pierrei
粘木
Ixonanthes
reticulata
海南油杉 Keteleeria hainanensis0.081
油丹 Alseodaphne hainanensis0.1840.144
海南韶子 Nephelium topengii0.6720.1030.656
土沉香 Aquilaria sinensis0.4960.2580.4360.546
普洱茶 Camellia sinensis var. assamica0.2450.8720.2590.2430.365
海南梧桐 Firmiana hainanensis0.3340.0000.0110.2230.3040.089
乐东拟单性木兰 Parakmeria lotungensis0.1100.2050.0690.0190.4850.2340.000
香子含笑 Michelia hedyosperma0.4200.0000.0020.1040.3720.2100.6930.000
陆均松 Dacrydium pierrei0.0590.0810.2610.1020.2250.0700.0000.4890.000
粘木 Ixonanthes reticulate0.2580.2850.2730.2090.3500.0190.0000.2250.0000.186
海南粗榧 Cephalotaxus mannii0.2820.0000.0080.2350.4310.1820.2850.0000.5890.0000.000

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3 讨论和结论

种群的生态位宽度是指一个种群在一个群落中所利用的不同资源的总和, 生态位宽度的大小取决于物种对环境的适应力、种间竞争强度和环境因子的分布状况。种群生态位宽度越大, 表明它对环境的适应能力越强, 对各种资源的利用越充分, 且往往在群落中处于优势地位(余世孝, 1995), 如对福建万木林珍稀植物的群落学研究表明, 生态位宽度大小与植物濒危原因没有必然的联系, 保护植物的生态位宽度可以是宽的, 也可以是窄的(毕晓丽, 2002); 对生态位宽度与环境因子梯度的研究(颜廷芬等, 1999)清楚地表明了各种群在不同环境下对资源的利用能力及对生态因子的适应能力, 进一步明确了各种群在群落中所处的地位以及群落的动态演替方向。同样, 在霸王岭保护区内, 一些物种的生态位宽度较大, 其分布范围也较广, 如: 土沉香、海南韶子、普洱茶等可以出现在许多群落中, 分布的海拔范围也较宽, 介于500-1400 m之间; 而一些物种, 如: 陆均松、鸡毛松为霸王岭山地雨林中的优势种, 但其海拔梯度上的分布范围狭窄, 仅集中分布在中高海拔的山地雨林中, 导致其生态位宽度较小, 进一步表明物种的生态位宽度不仅与其自身的生态适应力有关, 也与其自身的地理分布范围密切相关; 一些生长于特殊生境中的保护植物具有较大的生态位宽度, 如海南梧桐、海南粗榧往往出现在岩石裸露率较高、坡度较陡的生境中, 表明它们在特殊的生境中具有较强的适应能力, 也表明了它们在群落中具有重要的生态地位和作用。

许多研究证明, 生态位宽度较大的物种对资源的利用能力较强, 因而与其他种群间的生态位重叠也较大, 但是生态位宽度较大的物种的生物生态学特性不一定相同, 对资源的要求存在一定差异, 导致生态位宽度大的物种与其他物种的生态位重叠不一定大(王仁忠, 1997)。本研究同样表明, 一些生态位宽度较大的濒危植物, 如海南油杉与鸡毛松, 生态位重叠值仅为0.081; 而一些研究表明, 生态位宽度小的物种与其他物种的生态位重叠不一定小, 可能存在着两种原因: 一是物种间在共享资源的同时也存在着竞争关系; 二是植物对资源利用的同时也存在彼此促进的关系(张桂莲和张金屯, 2002)。本研究表明, 霸王岭保护区不同地形的小生境存在较大的差异, 导致生境的高度异质性, 小生境的复杂性和多样性致使植物与其生境特点相适应, 出现分布上的差异, 因此, 某些物种倾向于在局部适宜的生境斑块中具有较高的聚集度, 如: 香子含笑和粘木等的生态位宽度较小, 在坡度较大的沟谷中生长良好, 而在适宜生境斑块以外的空间分布较为贫乏, 以致这些物种的生态位宽度较小, 但有较高的生态位重叠。

物种之间的生态位重叠与种间竞争存在一定的关系, 在资源不足的情况下, 两个物种的生态位重叠在表征其生态相似性的同时, 还反映了它们之间存在的竞争关系, 而在资源丰富的情况下, 种间的生态位重叠并不反映其竞争程度(林思祖等, 2002), 只是表明物种之间因生态相似性占据了相似的生态空间。本研究中的一些保护植物, 如普洱茶与海南油杉在群落中的生态位重叠值最大, 为0.872, 但并非所有种均如此。因为群落的发展与演替是环境对物种自然选择的过程, 也是物种对环境的适应过程, 在环境压力和种间相互适应的双重作用下, 群落中的物种相互适应和协同进化, 在物种的形态和生理等方面出现了许多互补的特征, 使得同一群落中不同的物种利用资源的途径有所不同, 从而使物种达到与环境和谐与统一(张峰和上官铁梁, 2004; 赵永华等, 2004)。因此, 这可能是霸王岭濒危植物的生存群落中许多生态位重叠的物种能够共存, 形成相对稳定状态的一个原因。

综上所述, 有些保护物种, 如土沉香、海南韶子、普洱茶、油丹和海南油杉的生态位宽度较大, 其自身的生态适应力也较强, 这些物种主要受人为采挖和砍伐的影响, 对它们的保护重点是要减少人为破坏, 如有针对性地建设防护栏、巡护路、管护房、隔离带等必要的基础设施, 使其种群自然恢复; 而有些保护物种, 如香子含笑、粘木、海南梧桐和海南粗榧的生态位宽度较小, 个体数量较少, 仅分布在一些特殊的生境或群落中, 它们对环境的要求较苛刻, 在霸王岭所面临的威胁较严重, 对这些保护植物的研究与保护应该给予更多的关注, 除加强原生境的就地保护(如停止森林破坏、旅游建设)外, 还应开展这些植物的科学研究工作, 重点对它们的生长发育规律、生态适应性、遗传变异性, 以及如何提高它们对环境的适应幅度和引种栽培技术等方面进行研究。同时, 对这些物种的迁地保护还应充分考虑它们所适宜的生境及其占据的空间位置, 在资源不足的情况下, 这些物种比较容易被生态适应性较强的物种所排挤。另外, 陆均松、鸡毛松和乐东拟单性木兰虽为霸王岭山地雨林中的优势种, 但仅集中分布在中高海拔的局部地区, 地理分布狭窄, 对这些物种的保护应重点加强其所在的山地雨林群落的保护, 维持其群落的连贯性。因此, 未来对不同的濒危植物的保护, 不仅要考虑各物种自身的生物学特性, 还应从植物群落的角度出发。

致谢

感谢霸王岭国家级自然保护区管理局陈庆、王进强等护林员在野外调查中的帮助!

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Li DW, Zhang WH, Ren ZZ (2005).

Niche characteristics of dominant populations of Sophora davidii community in loess gully region

Chinese Journal of Applied Ecology, 16, 2231-2235. (in Chinese with English abstract)

URL     PMID:16515163      [本文引用: 2]

Based on field investigation data and employing Levin and niche proportional similarity formulae, this paper calculated the niche breadth, niche proportional similarity, and niche overlap of 13 dominant species of Sophora davidii community in loess gully region. The results showed that the niche breadth of the dominant populations of Sophora davidii community decreased in order of Sophora davidii (0.964), Artemisia giraldii (0.775), Arteniisia gmelinii (0.664), Thenmeda triandra var. japonica (0.508), Polygala tenuifolia (0.485), Setaria viridis (0.399), Cynodon dactylon (0.393), Bothriochloa ischaemum (0.327), Lespedeza davurica (0.321), Stipa bungeana (0.472), Periploca sepium (0.292), Clemnatis fruticosa (0.251), and Cleistogenes squarrosa (0.234). The niche proportional similarity between Sophora davidii and other species was 0.21-0.76, while the niche overlap was not high, which meant that there was no severe competition between them. It could be concluded that Sophora davidii was the dominant population in the community, with most resources utilization and most adaptability to the ecosystem.

[ 李登武, 张文辉, 任争争 (2005).

黄土沟壑区狼牙刺群落优势种群生态位研究

应用生态学报, 16, 2231-2235.]

URL     PMID:16515163      [本文引用: 2]

Based on field investigation data and employing Levin and niche proportional similarity formulae, this paper calculated the niche breadth, niche proportional similarity, and niche overlap of 13 dominant species of Sophora davidii community in loess gully region. The results showed that the niche breadth of the dominant populations of Sophora davidii community decreased in order of Sophora davidii (0.964), Artemisia giraldii (0.775), Arteniisia gmelinii (0.664), Thenmeda triandra var. japonica (0.508), Polygala tenuifolia (0.485), Setaria viridis (0.399), Cynodon dactylon (0.393), Bothriochloa ischaemum (0.327), Lespedeza davurica (0.321), Stipa bungeana (0.472), Periploca sepium (0.292), Clemnatis fruticosa (0.251), and Cleistogenes squarrosa (0.234). The niche proportional similarity between Sophora davidii and other species was 0.21-0.76, while the niche overlap was not high, which meant that there was no severe competition between them. It could be concluded that Sophora davidii was the dominant population in the community, with most resources utilization and most adaptability to the ecosystem.

Lin SZ, Huang SG, Hong W, Huang BL, Yu XT (2002).

The characteristics of multi-dimension niche of dominant populations in Chinese fir and broad-leaved mixed forest

Acta Ecologica Sinica, 22, 962-968. (in Chinese with English abstract)

[本文引用: 1]

[ 林思祖, 黄世国, 洪伟, 黄宝龙, 俞新妥 (2002).

杉阔混交林主要种群多维生态位特征

生态学报, 22, 962-968.]

[本文引用: 1]

Liu CS, Liu P, Zhang ZX, Li CH, Huang BW, Luo JF, Jin TT (2009).

The niche characteristics of endangered plant Tsuga chinensis var. tchekiangensis in Jiulongshan national nature reserve of Zhejiang Province

Journal of Wuhan Botanical Research, 27, 55-61. (in Chinese with English abstract)

[本文引用: 1]

[ 刘春生, 刘鹏, 张志祥, 李成惠, 黄帮文, 罗建峰, 金婷婷 (2009).

九龙山濒危植物南方铁杉的生态位研究

武汉植物学研究, 27, 55-61.]

[本文引用: 1]

Liu JF, Hong W (1999).

A study on the community ecology of Castanopsis kawakamii―study on the niche of the main tree population in Castanopsis kawakamii community

Acta Ecologica Sinica, 19, 347-352. (in Chinese with English abstract)

[本文引用: 1]

[ 刘金福, 洪伟 (1999).

格氏栲群落生态学研究: 格氏栲林主要种群生态位的研究

生态学报, 19, 347-352.]

[本文引用: 1]

Liu WD, Zang RG, Ding Y (2009).

Community features of two types of typical tropical monsoon forests in Bawangling Nature Reserve, Hainan Island

Acta Ecologica Sinica, 29, 3465-3476. (in Chinese with English abstract)

[本文引用: 1]

[ 刘万德, 臧润国, 丁易 (2009).

海南岛霸王岭两种典型热带季雨林群落特征

生态学报, 29, 3465-3476.]

[本文引用: 1]

Mueller LD, Altenberg L (1985).

Statistical inference on measures of niche overlap

Ecology, 66, 1204-1210.

DOI:10.2307/1939173      URL     [本文引用: 1]

Peng YS, Zheng MX, Mo LJ, Tang GD, Zhuang XY, Chen XM (2007).

Niches of dominant plant species of natural secondary forests in Zhuhai City, Guangdong Province

Chinese Journal of Ecology, 26, 483-488. (in Chinese with English abstract)

URL     [本文引用: 1]

A total of one hundred and twelve 10 m×10 m quadrats were set up in ten terrestrial
secondary forest communities of Zhuhai City, Guangdong Province to study the niche breadth and
niche overlap of the dominant plant species. The results indicated that the Shannon-Wiener’s niche
breadth and Levins’s niche breadth of 23 dominant species in the communities were 0.4049—0.7736
and 0.0551—0.3086, respectively. The species with larger niche breadth were Acronychia pedunculata,
Gordonia axillaris, Aporosa dioica, Microdesmis casearifolia, Gironniera subaequalis, and Sterculia lanceolata,
which were the widespread species in natural forests. The niche similarity of dominant species varied
between 0 and 0.3810, and the highest one occurred between A. dioica and Cinnamomum burmanii.
A. pedunculata had different degree niche overlaps with other 22 species, but G. axillaris only had niche
overlaps with a few dominant species. There were no niche overlaps in 20.6% of species-pairs. It was
suggested that some widespread and well-adapted species with larger niche breadth, e . g . , A. pedunculata,
G. axillaris
, G. subaequalis and Endospermum chinense, should be selected in prior in the restoration of
natural forests and the reconstruction of plantations.

[ 彭逸生, 郑明轩, 莫罗坚, 唐光大, 庄雪影, 陈锡沐 (2007).

珠海市陆生天然次生林优势种的生态位

生态学杂志, 26, 483-488.]

URL     [本文引用: 1]

A total of one hundred and twelve 10 m×10 m quadrats were set up in ten terrestrial
secondary forest communities of Zhuhai City, Guangdong Province to study the niche breadth and
niche overlap of the dominant plant species. The results indicated that the Shannon-Wiener’s niche
breadth and Levins’s niche breadth of 23 dominant species in the communities were 0.4049—0.7736
and 0.0551—0.3086, respectively. The species with larger niche breadth were Acronychia pedunculata,
Gordonia axillaris, Aporosa dioica, Microdesmis casearifolia, Gironniera subaequalis, and Sterculia lanceolata,
which were the widespread species in natural forests. The niche similarity of dominant species varied
between 0 and 0.3810, and the highest one occurred between A. dioica and Cinnamomum burmanii.
A. pedunculata had different degree niche overlaps with other 22 species, but G. axillaris only had niche
overlaps with a few dominant species. There were no niche overlaps in 20.6% of species-pairs. It was
suggested that some widespread and well-adapted species with larger niche breadth, e . g . , A. pedunculata,
G. axillaris
, G. subaequalis and Endospermum chinense, should be selected in prior in the restoration of
natural forests and the reconstruction of plantations.

Pianka ER (1973).

The structure of lizard communities

Annual Review of Ecology and Systematics, 4, 53-74.

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Robert M, Christoph N, Wilhelm B, Pierre LI (2003).

Biodiversity and endemism mapping as a tool for regional conservation planning―case study of the Pleurothallidinae (Orchidaceae) of the Andean rain forests in Bolivia

Biodiversity and Conservation, 12, 2005-2024.

DOI:10.1023/A:1024195412457      URL     [本文引用: 1]

Analyses of species diversity and endemism patterns provide vital inputs for conservation planning. Therefore, it is an important dilemma of biodiversity conservation that in very diverse but poorly studied tropical countries those patterns can hardly be considered. Consequently, there is an urgent need to develop prediction models that make the best use of existing data on species distribution and that can give hints on spatial conservation priorities. This paper presents the results of a pilot study on the diversity of the orchid subtribe Pleurothallidinae (331 mapped species) in the Andean rain forests of Bolivia. Results of a taxon-based mapping methodology, using abiotic (humidity and temperature, the latter indicated by altitude) and historical factors (taken into account as distance from collection localities) that determine species ranges, are compared with outcomes of an inventory-based mapping approach. The patterns of taxon-based diversity and endemism show a strong correlation with the distribution of sample localities. The inventory-based approach is more reliable, but it is interesting to apply both mapping methods in order to make a critical interpretation and comparison that facilitates some valuable conservation recommendations. We end with concrete conclusions for conservation planning and action.

Schoener TW (1970).

Non-synchronous spatial overlap of Lizards in patchy habitats

Ecology, 51, 408-418.

DOI:10.2307/1935376      URL     [本文引用: 1]

Shi HW, Jiang MX, Hu LL (2007).

Niche characteristics of Berchemiella wilsoni var. pubipetiolata, an endangered species in China

Journal of Wuhan Botanical Research, 25, 163-168. (in Chinese with English abstract)

[本文引用: 1]

[ 史红文, 江明喜, 胡理乐 (2007).

濒危植物毛柄小勾儿茶的生态位研究

武汉植物学研究, 25, 163-168.]

[本文引用: 1]

Shi XH, Xu XB, Zhang WH (2007).

Study on the niche of the main populations in Abies chinensis community

Bulletin of Botanical Research, 27, 345-349. (in Chinese with English abstract)

URL     [本文引用: 1]

Based on the data collected from the plots investigation along with the implementation Levins, Shannon weiner index and niche overlap formulas in Mt. Qinling Shaanxi, the niche breadth values, niche similarity and niche overlap values of the main tree species were measured in the forestry of Abies chensiensis in this study. The results showed that: with the respective Levins and Shannon weiner index, the results of niche breadths of the main populations were almost the same and the highest value of niche breadth in arbor layer, shrub layer and herb layer were Abies chensiensis, Sinarundinaria nitida and Carex lanceolata, respectively. Abies chensiensis was prone to form mixed forest with populations with high value of niche breadth. In the populations with high value of niche breadth, the niche similarity was generally higher accordingly. Niche overlaps extents of main populations in Abies chensiensis community were correspondingly lower, indicated that the environmental resource could be shared by every population and the community was comparative stable.

[ 史小华, 许晓波, 张文辉 (2007).

秦岭冷杉群落主要种群生态位研究

植物研究, 27, 345-349.]

URL     [本文引用: 1]

Based on the data collected from the plots investigation along with the implementation Levins, Shannon weiner index and niche overlap formulas in Mt. Qinling Shaanxi, the niche breadth values, niche similarity and niche overlap values of the main tree species were measured in the forestry of Abies chensiensis in this study. The results showed that: with the respective Levins and Shannon weiner index, the results of niche breadths of the main populations were almost the same and the highest value of niche breadth in arbor layer, shrub layer and herb layer were Abies chensiensis, Sinarundinaria nitida and Carex lanceolata, respectively. Abies chensiensis was prone to form mixed forest with populations with high value of niche breadth. In the populations with high value of niche breadth, the niche similarity was generally higher accordingly. Niche overlaps extents of main populations in Abies chensiensis community were correspondingly lower, indicated that the environmental resource could be shared by every population and the community was comparative stable.

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[本文引用: 1]

Wang G, Zhao SL, Zhang PY, Chen QC (1984).

Study on the definition of niche and the improved formula for measuring niche overlap

Acta Ecologica Sinica, 4, 119-127. (in Chinese with English abstract)

[本文引用: 2]

[ 王刚, 赵松岭, 张鹏云, 陈庆诚 (1984).

关于生态位定义的探讨及生态位重叠计测公式改进的研究

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[本文引用: 2]

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Age structure and niche of the endangered Magnolia sieboldii in Huangshan Mountain

Acta Ecologica Sinica, 26, 1862-1871. (in Chinese with English abstract)

[本文引用: 1]

[ 王立龙, 王广林, 黄永杰, 李晶, 刘登义 (2007).

黄山濒危植物小花木兰生态位与年龄结构研究

生态学报, 26, 1862-1871.]

[本文引用: 1]

Wang RZ (1997).

The niche breadths and niche overlaps of main plant populations in Leymus chinensis grassland for grazing

Acta Phytoecologica Sinica, 21, 304-311. (in Chinese with English abstract)

[本文引用: 1]

[ 王仁忠 (1997).

放牧影响下羊草草地主要植物种群生态位宽度与生态位重叠的研究

植物生态学报, 21, 304-311.]

[本文引用: 1]

Wang ZN, He KN, Zhang WQ, Liu CF, Liu S (2005).

Study on niche characteristics of undergrowth vegetation during plant restoration in semi-arid region

Journal of Soil and Water Conservation, 19(5), 162-165. (in Chinese with English abstract)

DOI:10.5958/2455-7145.2020.00022.3      URL     [本文引用: 1]

[ 王正宁, 贺康宁, 张卫强, 刘晨峰, 刘胜 (2005).

半干旱地区植被恢复过程中林下植被生态位特征的研究

水土保持学报, 19(5), 162-165.]

DOI:10.5958/2455-7145.2020.00022.3      URL     [本文引用: 1]

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Measuring realized niche spaces climatic response of chaparral and coastal sage scrub

Ecology, 72, 1678-1684.

DOI:10.2307/1940967      URL     [本文引用: 1]

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Effect analysis of environmental factors on plant niche breadth

Journal of Northeast Forestry University, 27, 35-38. (in Chinese with English abstract)

[本文引用: 1]

[ 颜廷芬, 丛沛桐, 刘兴华, 祖元刚 (1999).

环境因子对植物生态位宽度影响程度分析,

东北林业大学学报, 27, 35-38.]

[本文引用: 1]

Yu SX (1995). Introduction to Mathematical Ecology. Scientific and Technical Documentation Press, Beijing. (in Chinese)

[本文引用: 1]

[ 余世孝 (1995). 数学生态学导论. 科学技术文献出版社, 北京.]

[本文引用: 1]

Zang RG, Yang YC, Lin RC, Yang XS (2003).

Study on the forest cycle and community characteristics in a tropical montane rain forest in Bawangling, Hainan Province

Scientia Silvae Sinicae, 39(5), 1-9. (in Chinese with English abstract)

DOI:10.11707/j.1001-7488.20030501      URL     [本文引用: 1]

According to the theory of forest growth cycle, forest communities are dynamic mosaic systems composed of patches in different developmental stages. On basis of the quandrats investigation carried out in a 1 hm2 tropical montane rain forest in Bawangling, Hainan Province, the community characteristics of the whole forest stand were analyzed.The population and community features were studied based on the theory of forest cycle. The result showed that the utilization of ecological resources in the horizontal and vertical directions for trees of different life forms were relatively separate. Most shrubs and small statured trees had greater niche breadth values in the horizontal direction, and had less niche breadth values in the vertical direction ,and those for the middle or tall trees were on the contrary. The niche breadth for the rare and low density species were the least in both directions. With the progress of forest cycle, the frequency of different species presented in different patch types changed accordingly. The distribution patterns for most dominant trees and shrubs in the community were clumped, those for the trees having moderate individuals were uniform, and those for the low density species were random. In the process of forest cycle, the individuals owned by each species and the space occupied by each species decreased gradually. The numbers of species in different patch types in the forest cycle decreased with the increase of numbers of individuals. The numbers of species with 1 individuals and 2 individuals were most abundant ,and the numbers of species with more than 3 individuals decreased apparently in the investigated forest. Similarity analysis showed that the dynamic changes of patches in the forest cycle were somewhat continuous and gradual, the similarities between the neighboring stages were greater than those non_neighboring stages in the forest cycle. Calculation of niche breadth for different species on basis of the theory of forest cycle showed that the utilization of ecological resources in different patches by the low density species or those confined only in certain patches were comparatively less, while those species presented in most patches and had moderate numbers of individuals or moderate basal areas had greater utilizations for the ecological resources in the forest cycle.

[ 臧润国, 杨彦承, 林瑞昌, 杨秀森 (2003).

海南霸王岭热带山地雨林森林循环与群落特征研究

林业科学, 39(5), 1-9.]

DOI:10.11707/j.1001-7488.20030501      URL     [本文引用: 1]

According to the theory of forest growth cycle, forest communities are dynamic mosaic systems composed of patches in different developmental stages. On basis of the quandrats investigation carried out in a 1 hm2 tropical montane rain forest in Bawangling, Hainan Province, the community characteristics of the whole forest stand were analyzed.The population and community features were studied based on the theory of forest cycle. The result showed that the utilization of ecological resources in the horizontal and vertical directions for trees of different life forms were relatively separate. Most shrubs and small statured trees had greater niche breadth values in the horizontal direction, and had less niche breadth values in the vertical direction ,and those for the middle or tall trees were on the contrary. The niche breadth for the rare and low density species were the least in both directions. With the progress of forest cycle, the frequency of different species presented in different patch types changed accordingly. The distribution patterns for most dominant trees and shrubs in the community were clumped, those for the trees having moderate individuals were uniform, and those for the low density species were random. In the process of forest cycle, the individuals owned by each species and the space occupied by each species decreased gradually. The numbers of species in different patch types in the forest cycle decreased with the increase of numbers of individuals. The numbers of species with 1 individuals and 2 individuals were most abundant ,and the numbers of species with more than 3 individuals decreased apparently in the investigated forest. Similarity analysis showed that the dynamic changes of patches in the forest cycle were somewhat continuous and gradual, the similarities between the neighboring stages were greater than those non_neighboring stages in the forest cycle. Calculation of niche breadth for different species on basis of the theory of forest cycle showed that the utilization of ecological resources in different patches by the low density species or those confined only in certain patches were comparatively less, while those species presented in most patches and had moderate numbers of individuals or moderate basal areas had greater utilizations for the ecological resources in the forest cycle.

Zang RG, Yang YC, Jiang YX (2001).

Community structure and tree species diversity characteristics in a tropical montane rain forest in Bawangling Nature Reserve, Hainan Island

Acta Phytoecologica Sinica, 25, 270-275. (in Chinese with English abstract)

[本文引用: 1]

[ 臧润国, 杨彦承, 蒋有绪 (2001).

海南岛霸王岭热带山地雨林群落结构及树种多样性特征的研究

植物生态学报, 25, 270-275.]

[本文引用: 1]

Zhang F, Shangguan TL (2004).

Niche characteristics of dominant populations in Elaeagnus mollis communities, Shanxi

Acta Botanica Boreali-Occidentalia Sinica, 24, 70-74. (in Chinese with English abstract)

[本文引用: 1]

[ 张峰, 上官铁梁 (2004).

翅果油树群落优势种群生态位分析

西北植物学报, 24, 70-74.]

[本文引用: 1]

Zhang GL, Zhang JT (2002).

Niche analysis of dominant species in Shenweigou of Guandi Mountain

Journal of Wuhan Botanical Research, 20, 203-208. (in Chinese with English abstract)

[本文引用: 1]

[ 张桂莲, 张金屯 (2002).

关帝山神尾沟优势种群生态位分析

武汉植物学研究, 20, 203-208.]

[本文引用: 1]

Zhang JT (2004). Quantitative Ecology. Science Press, Beijing. (in Chinese)

[本文引用: 1]

[ 张金屯 (2004). 数量生态学. 科学出版社, 北京.]

[本文引用: 1]

Zhang ZD, Zang RG (2007).

Influence of ecological factors on distribution of woody plant functional types in a natural tropical forest landscape, Bawangling, Hainan Island, South China

Journal of Plant Ecology (Chinese Version), 31, 1092-1102. (in Chinese with English abstract)

[本文引用: 2]

[ 张志东, 臧润国 (2007).

海南岛霸王岭热带天然林景观中木本植物功能型分布的影响因素

植物生态学报, 31, 1092-1102.]

[本文引用: 2]

Zhao YH, Lei RD, He XY, Jia X (2004).

Niche characteristics of plant populations in Quercus aliena var. acuteserrata stands in Qinling Mountains

Chinese Journal of Applied Ecology, 15, 913-918. (in Chinese with English abstract)

URL     PMID:15362607      [本文引用: 1]

Based on the data collected from 31 plots and using Levins, Hurlbert and Pianka formulas, this paper calculated and analyzed the niche breadths and overlaps of 24 tree and 29 shrub populations in Quercus aliena var. acuteserrata stands in Mt. Qinling, Shaanxi. The results showed that the Levins and Hurlbert's niche breadths of Quercus aliena var. acuteserrata , Pinus armandi, Toxicodendron vernicifluum, Populus davidiana and Pinus tabulaeformis in arbor layer were 3.695, 1.695, 1.325, 0.840, 0.702 and 0.036, 0.299, 0.568, 1.721, 2.701 respectively, and those of Corrylus heterophylla, Lespedeza formosa and Smilax vaginata in shrub layer were 1.833, 1.466, 0.984 and 0.111, 0.300, 0.594, respectively. In general, the niche overlaps between the species with bigger niche breadth and other species were higher, and those of the species having similar environmental requirements were also higher.

[ 赵永华, 雷瑞德, 何兴元, 贾夏 (2004).

秦岭锐齿栎林种群生态位特征研究

应用生态学报, 15, 913-918.]

URL     PMID:15362607      [本文引用: 1]

Based on the data collected from 31 plots and using Levins, Hurlbert and Pianka formulas, this paper calculated and analyzed the niche breadths and overlaps of 24 tree and 29 shrub populations in Quercus aliena var. acuteserrata stands in Mt. Qinling, Shaanxi. The results showed that the Levins and Hurlbert's niche breadths of Quercus aliena var. acuteserrata , Pinus armandi, Toxicodendron vernicifluum, Populus davidiana and Pinus tabulaeformis in arbor layer were 3.695, 1.695, 1.325, 0.840, 0.702 and 0.036, 0.299, 0.568, 1.721, 2.701 respectively, and those of Corrylus heterophylla, Lespedeza formosa and Smilax vaginata in shrub layer were 1.833, 1.466, 0.984 and 0.111, 0.300, 0.594, respectively. In general, the niche overlaps between the species with bigger niche breadth and other species were higher, and those of the species having similar environmental requirements were also higher.

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