自然保护区旅游开发对不同扩散模式植物多样性的影响
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摘要
作为一种强烈的干扰源,旅游开发活动已经明显地改变了世界上的许多景观,导致了大量的生态和环境问题,对生物多样性产生了严重威胁。通常,旅游开发对生物多样性的影响起始于游憩带的边缘效应。为了深入探讨这些边缘效应,以景观生态学为理论基础,对旅游景观生态系统的空间要素、结构和功能特征进行了分析和讨论,然后以植物个体的生活史特征——种子扩散和植物运动生态学框架为依据建立研究假设,来检验旅游活动对不同扩散模式植物多样性的影响机制。
实证研究选取兴凯湖自然保护区湖岗游憩带作为研究对象,实验起止时间为2006至2012年。整个实验分为两个部分,第一部分是对游憩带(包括旅游斑块和交通廊道)具有不同扩散模式的植物(动物扩散与风扩散物种)多样性的时空动态进行检验。方法是分别选择不同干扰强度的游憩带,对所设置样方内的所有维管植物进行记录,并将它们分为动物扩散(animal-dispersed)和风扩散(wind-dispersed)两个群体,通过对它们物种多样性动态的检验来反映旅游开发对游憩带植物扩散的影响；第二阶段是对第一部分实验的补充,通过对干扰带和非干扰带昆虫传粉和鸟类扩散种实行为差异性的比较来进一步验证旅游开发对动物扩散物种的影响机制。结果显示：
(1)在过去的6年间,两个实验景观中的动物扩散物种和风扩散物种表现了明显不同的边缘效应：强干扰带的动物扩散物种在所有距离序列上均有不同程度的减少,而弱干扰带在10-30m的距离序列上不但没有减少相反有所增加；风扩散物种虽然在两个实验景观中均有所增加,但在强干扰带表现了明显的边缘效应,而在弱干扰带,则表现了明显的连通效应。两者的动态最终导致了总体多样性的变化；
(2)游憩带的访花昆虫在物种多样性、访花频率和停留时间等方面均显著低于非游憩带；观察发现,游人干扰是造成访花时间变短的主要原因,而游憩带访花昆虫丰富度与访花频度显著低于非游憩带可能是景观破碎化和游人干扰共同导致的结果；
(3)旅游活动(游人活动和交通干扰)对食果鸟的取食行为产生了显著影响,减少了幼苗更新数量。这一方面是因为旅游干扰影响了鸟类的取食时间,另一方面也可能是生境破碎化导致了取食鸟的丰富度与取食频度的减少。此外,那些未经完全取食而直接落于母株树冠下的种实也可能由于母株的自毒作用或竞争性死亡而减少幼苗自然更新的数量；
(4)游人活动干扰并未对幼苗更新格局产生显著影响,但交通干扰却对幼苗的分布格局产生了明显影响：幼苗分布格局明显偏向于斑块内部,而非干扰带则明显偏向于交通廊道一侧。
检验结果表明,旅游开发对动物扩散物种和风扩散物种产生了明显影响,导致了不同的边缘效应。这可能来自于两种机制：(1)旅游开发改变了景观连通度,影响了不同扩散媒介(如鸟和风)对种子的扩散；(2)游憩活动通过干扰动植物的互利关系(传粉和种子传播)对种子扩散产生影响。这充分证明了旅游开发对植物多样性影响的复杂性：它不仅可以通过降低生境质量产生影响,也可能通过干扰生态过程如种间关系对其产生连锁影响。这种由旅游开发而导致的“渗透干扰效应”,可能对景观内某些动物扩散物种的长期存续具有更大的威胁性。该研究模式为旅游干扰研究提供了新的视角。
As a type of interference, tourism developments have considerably changed many landscapes of the world, resulting in a large number of ecological and environmental issues, threatening to biodiversity. The impacts of tourism developments on biodiversity are usually resulted from edge-effects. In order to further investigate this edge-effect, the paper analyzed and discussed the characters of tourism landscape ecosystem in aspects of spatial constructions and functions based on the theory of landscape ecology, and then test the disturbance mechanisms of tourism activities to biodiversity of plants dispersed by various vectors by establishing research hypothesis based on plant seed-dispersal and plant individual emotion ecology framework.
The study was conducted in Khanka Nature Reserve from2006to2012, and two recreation landscapes exerted different disturbance intensity were selected within a lakefront forest. Across experimental years, all vascular plants recorded were divided into animal-dispersed species and wind-dispersed species. The purpose was to present the impacts of tourism development on plant motion ecology by monitoring the dynamics of plant diversity at spatial and temporal scales. To further test the disturbance mechanisms of tourism activities to animal-dispersed species, the experiments were also conducted on pollination and seed dispersal. The results showed that:
(1) Across6years, animal-dispersed species and wind-dispersed species in the two experimental landscapes presented considerably different edge-effects:animal-dispersed species decreased across distances in both HDRP and LDRP, while wind-dispersed species all increased in the two landscapes. HDRP showed clear edge-effects while LDRP showed clear connectivity-effects in species richness. The dynamics of these two groups led to the changes of total diversity in the two landscapes.
(2) Recreation ecotone was significantly lower than non-recreational zone in species richness of pollinators, visiting frequency and time; visitor disturbances mainly led to the reduction of visiting time; Landscape fragmentation and tourist disturbances might jointly led to the reductions in species richness and visiting frequency of pollinators;
(3) Tourism activities (visitors and traffic disturbances) affected the behaviors of frugivorous birds, leading to the reductions of recruits. This might also be attributed to two mechanisms: tourism disturbances and landscape isolation. In addition, some fruits fell under mother plant canopy might be killed by toxicity of mother plants or competition between individuals, and thus decreased the numbers of recruits;
(4) Visitor activities did not considerably affect the distributions of seedlings, but traffic disturbances exerted significant effects on it.
The results indicate that, tourism development can exert different effects on plants dispersed by different vectors, which may be attributed to two mechanisms:the disruptions of both landscape modification and visitor's activities to the mutualisms of animal-plant. This demonstrates that tourism development can take ripple effects on plants by affecting some animals. The Halo disturbance from patch-edges may considerably threaten to the species dispersed by animals. The study provides new insight into tourism disturbances.
实证研究选取兴凯湖自然保护区湖岗游憩带作为研究对象,实验起止时间为2006至2012年。整个实验分为两个部分,第一部分是对游憩带(包括旅游斑块和交通廊道)具有不同扩散模式的植物(动物扩散与风扩散物种)多样性的时空动态进行检验。方法是分别选择不同干扰强度的游憩带,对所设置样方内的所有维管植物进行记录,并将它们分为动物扩散(animal-dispersed)和风扩散(wind-dispersed)两个群体,通过对它们物种多样性动态的检验来反映旅游开发对游憩带植物扩散的影响；第二阶段是对第一部分实验的补充,通过对干扰带和非干扰带昆虫传粉和鸟类扩散种实行为差异性的比较来进一步验证旅游开发对动物扩散物种的影响机制。结果显示：
(1)在过去的6年间,两个实验景观中的动物扩散物种和风扩散物种表现了明显不同的边缘效应：强干扰带的动物扩散物种在所有距离序列上均有不同程度的减少,而弱干扰带在10-30m的距离序列上不但没有减少相反有所增加；风扩散物种虽然在两个实验景观中均有所增加,但在强干扰带表现了明显的边缘效应,而在弱干扰带,则表现了明显的连通效应。两者的动态最终导致了总体多样性的变化；
(2)游憩带的访花昆虫在物种多样性、访花频率和停留时间等方面均显著低于非游憩带；观察发现,游人干扰是造成访花时间变短的主要原因,而游憩带访花昆虫丰富度与访花频度显著低于非游憩带可能是景观破碎化和游人干扰共同导致的结果；
(3)旅游活动(游人活动和交通干扰)对食果鸟的取食行为产生了显著影响,减少了幼苗更新数量。这一方面是因为旅游干扰影响了鸟类的取食时间,另一方面也可能是生境破碎化导致了取食鸟的丰富度与取食频度的减少。此外,那些未经完全取食而直接落于母株树冠下的种实也可能由于母株的自毒作用或竞争性死亡而减少幼苗自然更新的数量；
(4)游人活动干扰并未对幼苗更新格局产生显著影响,但交通干扰却对幼苗的分布格局产生了明显影响：幼苗分布格局明显偏向于斑块内部,而非干扰带则明显偏向于交通廊道一侧。
检验结果表明,旅游开发对动物扩散物种和风扩散物种产生了明显影响,导致了不同的边缘效应。这可能来自于两种机制：(1)旅游开发改变了景观连通度,影响了不同扩散媒介(如鸟和风)对种子的扩散；(2)游憩活动通过干扰动植物的互利关系(传粉和种子传播)对种子扩散产生影响。这充分证明了旅游开发对植物多样性影响的复杂性：它不仅可以通过降低生境质量产生影响,也可能通过干扰生态过程如种间关系对其产生连锁影响。这种由旅游开发而导致的“渗透干扰效应”,可能对景观内某些动物扩散物种的长期存续具有更大的威胁性。该研究模式为旅游干扰研究提供了新的视角。
As a type of interference, tourism developments have considerably changed many landscapes of the world, resulting in a large number of ecological and environmental issues, threatening to biodiversity. The impacts of tourism developments on biodiversity are usually resulted from edge-effects. In order to further investigate this edge-effect, the paper analyzed and discussed the characters of tourism landscape ecosystem in aspects of spatial constructions and functions based on the theory of landscape ecology, and then test the disturbance mechanisms of tourism activities to biodiversity of plants dispersed by various vectors by establishing research hypothesis based on plant seed-dispersal and plant individual emotion ecology framework.
The study was conducted in Khanka Nature Reserve from2006to2012, and two recreation landscapes exerted different disturbance intensity were selected within a lakefront forest. Across experimental years, all vascular plants recorded were divided into animal-dispersed species and wind-dispersed species. The purpose was to present the impacts of tourism development on plant motion ecology by monitoring the dynamics of plant diversity at spatial and temporal scales. To further test the disturbance mechanisms of tourism activities to animal-dispersed species, the experiments were also conducted on pollination and seed dispersal. The results showed that:
(1) Across6years, animal-dispersed species and wind-dispersed species in the two experimental landscapes presented considerably different edge-effects:animal-dispersed species decreased across distances in both HDRP and LDRP, while wind-dispersed species all increased in the two landscapes. HDRP showed clear edge-effects while LDRP showed clear connectivity-effects in species richness. The dynamics of these two groups led to the changes of total diversity in the two landscapes.
(2) Recreation ecotone was significantly lower than non-recreational zone in species richness of pollinators, visiting frequency and time; visitor disturbances mainly led to the reduction of visiting time; Landscape fragmentation and tourist disturbances might jointly led to the reductions in species richness and visiting frequency of pollinators;
(3) Tourism activities (visitors and traffic disturbances) affected the behaviors of frugivorous birds, leading to the reductions of recruits. This might also be attributed to two mechanisms: tourism disturbances and landscape isolation. In addition, some fruits fell under mother plant canopy might be killed by toxicity of mother plants or competition between individuals, and thus decreased the numbers of recruits;
(4) Visitor activities did not considerably affect the distributions of seedlings, but traffic disturbances exerted significant effects on it.
The results indicate that, tourism development can exert different effects on plants dispersed by different vectors, which may be attributed to two mechanisms:the disruptions of both landscape modification and visitor's activities to the mutualisms of animal-plant. This demonstrates that tourism development can take ripple effects on plants by affecting some animals. The Halo disturbance from patch-edges may considerably threaten to the species dispersed by animals. The study provides new insight into tourism disturbances.
引文
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