食果动物传播植物种子的有效性

传播有效性是评估食果动物对植物种群更新贡献的关键指标,它决定了植物种群的续存,影响着群落生物多样性的维系。本文从种子传播数量、质量角度分析了影响动物传播有效性的因素,并从有效传播动物组合角度分析了其对植物种群更新的影响。传播数量的多少与动物的取食行为有关。以整吞取食的食果动物常被认为是植物的有效传播动物,而植物的果实为动物偏好选择的食物时,势必导致该种植物拥有高的种子传播数量。传播质量与食果动物肠道种子滞留时间以及其后的种子传播距离和排放地相关。体型较大、生境适应能力和迁徙性较强的动物常为植物提供较远的传播距离,而被认为传播质量较高。取食后动物的生境选择决定了种子的排放地,进而影响它们的种子传播质量。如果取食后动物偏好利用植物适宜更新的生境时,动物则拥有较高的传播质量。有效传播动物通常在传播数量和传播质量以互补形成一些组合对植物种群更新起贡献。今后工作应将动物行为野外监测和种子萌发实验相结合、取食后多种行为相结合来分析动物的传播有效性;而以一些国内特有濒危植物为研究对象开展传播有效性研究应是今后恢复生态学研究的重点。     

植物种群更新限制——从种子生产到幼树建成

更新限制是指种子由于各种原因,不能够萌发并生长成幼树。它作为解释生物多样性的理论,一直受到国内外群落生态学家关注。从种源限制、传播限制和建成限制3个角度,对更新限制机制研究进展进行了综述。从种源限制而言,时空因素是影响植物种群更新限制的重要因素,因为植物结实量存在明显时空变化,造成植物更新个体出现明显的时空规律。从传播限制而言,传播数量、距离和食果动物行为均限制植物种群更新。数量上,缺乏有限传播者势必减少传播数量,但如果种子拥有较高质量,则能逃脱数量限制;距离上,植物更新个体显示出明显的Janzen-Connell格局,但传播距离趋向稳定,形成植物种群的进化稳定对策;食果动物行为上,不同传播者对更新贡献存在差异,捕食者直接降低更新,融入两类动物行为的模型更能反映食果动物对更新的限制。从建成限制而言,环境因子制约植物生长。小尺度下,微生境的好坏对于植物幼苗建成至关重要;大尺度下,植物提供较好的广告效应则能摆脱生境限制。将传播者行为、捕食者行为与幼苗的空间分布格局、种子传播机理模型等结合,建立植物更新限制机理模型应是更新限制未来的研究热点。选择稀有种和古老种为主题的长期更新限制研究,为种群恢复提供指导,也是未来重要研究方向。     

生境破碎化对食果动物及种子传播的影响

食果动物与依赖其传播种子的植物间在进化过程中形成互惠关系,生境破碎化往往干扰种子传播过程,继而破坏这种关系.生境破碎化通常降低食果动物的多样性,但亦有相反的情况出现.食果动物对生境破碎化的适应能力不同,泛性森林动物和广食性动物具有较强的适应性.生境破碎化对依赖动物传播的植物影响有差异,多数植物受到负面影响,但也有一些植物不受影响,甚至受益.动物在破碎生境中对种子传播的有效性是种子搬运量、传播距离、种子萌发及种群建立等环节的综合效果.破碎化生境中种子的搬运量与动物的觅食行为和食物选择有关;种子传播距离受食物资源可获得性的改变和生境斑块异质性的影响;种子萌发和更新种群建立成功与否决定于是否存在有效的种子传播者.生境破碎化如何影响种子传播以及动植物相互关系,尤其是异质斑块的空间分布如何影响食果动物的传播有效性、破碎化生境下动植物互惠共生关系如何建立,生境破碎化导致的植物入侵对本地植物种子传播的影响是未来需要深入研究的问题.     

动物分散贮食行为对植物种群更新的影响

分散贮食是许多动物取食行为策略的重要组成部分。对以植物种子为主要贮食对象的动物来说,种子内营养物质含量、种子大小以及种子内次生化合物的含量等因素都直接影响动物的贮食行为。动物偏爱贮藏个体较大的种子,大种子多被搬运并分散贮藏在远离种源的地方,而小种子则多被就地取食,以补充动物贮食过程中的能量消耗。贮食动物主要通过空间记忆、特殊路线以及贮藏点周围的直接线索等途径重新获取贮藏点内食物。在重取过程中,一些贮藏点被遗忘,其中的种子成为植物种群更新的潜在种子库。因此,分散贮食动物不仅是种子捕食者,还是种子传播者,它们对植物种子的捕食、搬运和贮藏,影响了植物种子的存活和幼苗的建成,从而在一定程度上影响植物种群的更新、分布。植物种群为了促进种子的传播,在进化过程中逐渐形成了形式多样的适应性策略,降低种子的直接被捕食率,提高种子的被贮藏率。研究动物分散贮食行为对植物种群更新的影响,将有助于理解贮食动物与植物之间的互惠关系,从而认识贮食动物种群在生态系统中的作用,为生物多样性的保护提供科学依据。     

云南拉沙山滇金丝猴的种子传播

灵长类是森林中重要的种子传播者,其行为对于森林植物的更新具有重要的生态意义.虽然大量国外科学家关注了灵长类的种子传播行为,但我国的灵长类种子传播研究依然处于起步阶段,这不利于厘清我国灵长类的种子传播行为特点.本研究以云南拉沙山为研究地,关注滇金丝猴对果实性植物的取食及其对种子的传播作用,通过在夜栖地和午休地收集粪便分析灵长类的种子传播特点.结果表明:滇金丝猴共取食9科14种果实性植物,其中蔷薇科植物相对较多,共有6种,占所有植物种类的42.86%.果实特征影响着滇金丝猴的食物选择,滇金丝猴偏好选择肉质果植物和红色果实植物,分别有12种和8种,各占总数的85.71%和57.14%.不同类型的栖息地中,滇金丝猴粪便排放的种子有所差异,75%的种子被排放至夜栖地,仅有25%被排放至午休地.在夜栖地中,大量完整的蔷薇科植物种子被滇金丝猴排放至冷杉林中,这对于某些阔叶树种子在冷杉林中更新至关重要.本研究初步证实了滇金丝猴作为种子传播者在生态系统植物更新中的重要作用.     

斑块生境中食果鸟类对南方红豆杉种子的取食和传播

生境斑块化对食果鸟类的移动行为产生影响,进而影响其对植物种子的传播格局和效率。南方红豆杉(Taxus chinensis var.mairei)是我国一级保护植物,在南方山区多因人为干扰而呈斑块状分布。2011年,2012年10月底到12月中,以分布于福建梅花山国家级自然保护区红豆杉生态园中南方红豆杉种群为对象,研究斑块生境中鸟类对南方红豆杉种子的取食和传播行为,并评估专性鸟类和泛性鸟类的传播效率。结果发现:南方红豆杉源斑块共吸引22种食果鸟类取食红豆杉种子,并与13种鸟类形成了种子传播关系。不同年间,黑短脚鹎(Hypsipetes leucocephalus)都是植物的主要传播鸟类,而其他鸟类传播者种类具有一定的年间变化。生境斑块化导致专性鸟类黑短脚鹎和泛性鸟类红嘴蓝鹊(Urocissa erythrorhyncha)种子传播效率差异。与红嘴蓝鹊相比,黑短脚鹎飞行的平均距离较短((16.3±11.0)m,Mean±SD,n=125),传播距离相对较近;且它们取食后偏好在源斑块中活动,喜栖息在同种成树、甜楮(Castanopsis eyrei)及其他阔叶树等栖树上。红嘴蓝鹊取食后常在斑块间移动,常停歇在同种成树和毛竹(Phyllostachys heterocycla)上,传播距离相对较远((24.9±20.0)m,Mean±SD,n=95)。空间一致性结果表明,黑短脚鹎移动距离对幼苗更新距离的空间一致性程度高;而红嘴蓝鹊偏好生境与幼苗更新生境一致性程度高。结果表明,斑块生境中植物能与专性鸟类、泛性鸟类之间形成种子传播互惠关系,且种子传播效率受到专性鸟类和泛性鸟类传播距离和传播生境的影响。     

干扰对动物传播森林植物种子有效性影响的研究进展

干扰在森林生态系统中普遍存在,并影响森林的更新和演替.动物传播种子是种子更新的必经阶段,其对森林干扰的响应在一定程度上能够预测未来的森林群落组成和结构变化,对于明确森林演替方向具有重要意义.本文论述了森林干扰对动物传播种子有效性(包括动物传播种子的数量和质量)影响研究的生态学意义,全面揭示了自然干扰(火干扰、林窗干扰等)和人为干扰(生境破碎化、狩猎、采伐等)对动物传播种子数量、传播距离以及传播后幼苗更新影响的研究进展,指出干扰通过影响动物种群动态,进而造成动物传播种子数量发生了改变,动物传播种子的距离对干扰的响应基本表现出轻微负相关;干扰对传播后幼苗更新的影响结果因干扰类型的不同而复杂多变,干扰迹地环境因子的变化也影响着传播后的种子萌发和幼苗更新.干扰对动物传播种子有效性影响研究中存在的问题,主要表现为火干扰迹地恢复过程、增益性的干扰(如抚育、间伐、林窗)等对种子传播有效性影响研究的匮乏,以及忽略了温带森林内的干扰对动物传播种子的影响等.今后,应开展干扰对种子传播有效性的长期研究;对于干扰多发地带的森林,应高度重视增益性干扰影响动物传播植物种子的研究.     

高黎贡山天行长臂猿的果实性食物组成及种子传播潜力

灵长类动物是森林植物种子的重要传播媒介,它们的取食行为和栖息地利用影响着森林植物的种群更新.天行长臂猿是云南高黎贡山的明星物种之一.研究天行长臂猿的果实性食物和种子传播潜力有助于认清濒危动物的种子传播功能并评估它们对森林植物更新的贡献.于2017年10月-2018年12月,对云南高黎贡山赧亢的天行长臂猿家群的果实性食物...     

种子大小及其命运对植被更新贡献研究进展

种子是种子植物的繁殖体.种子对后代的资源投入将会直接影响后代的适合度进而影响到植被群落的更新.一般较大种子物种在种子萌发和幼苗生长阶段是具有较大优势的;但较小种子物种在逃避被动物采食和形成持久的土壤种子库成为植被更新的后备动力方面具有较大的优势.在不同的选择压力下,不同大小种子在以后的生活史中具有不同的命运,对植被的更新也具有不同的贡献.本文主要从种子大小和种子的传播散布、种子萌发、种子存活以及种子土壤库等方面综述了国外关于种子大小及其以后命运对植被更新贡献的研究结果,并对国内的相关研究提出了一些建议：种子生态学研究需要结合群落特点以及微环境等,开展从种子生产到幼苗更新全过程的系统性研究,为植物物种多样性保育和退化植被恢复方面提出有科学依据的管理措施.     

灵长类动物对植物种子的传播作用

种子传播对植物种群空间分布和群落生物多样性维系有着深远的影响。森林灵长类动物可有效地传播植物种子,其对果实的选择不仅受其自身形态、生理和行为特征的影响,也与果实的形态特征和同域分布的其他动物类群的取食竞争密切相关。灵长类动物传播种子的模式可分为消化道传播、携带传播和颊囊传播,其中消化道传播距离远、种子萌发或幼苗建成率高,是最为有效的传播模式。灵长类动物对种子的传播作用还能促进退化生境的恢复。建议今后研究中应考虑灵长类动物的多途径种子传播和多种因素对种子扩散与萌发的综合影响,同时要更加客观地评估灵长类动物缺失对森林植被更新的影响。     

Seed-deposition and recruitment patterns of Clusia species in a disturbed tropical montane forest in Bolivia

Spatial patterns of seed dispersal and recruitment of fleshy-fruited plants in tropical forests are supposed to be driven by the activity of animal seed dispersers, but the spatial patterns of seed dispersal, seedlings and saplings have rarely been analyzed simultaneously. We studied seed deposition and recruitment patterns of three Clusia species in a tropical montane forest of the Bolivian Andes and tested whether these patterns changed between habitat types (forest edge vs. forest interior), distance to the fruiting tree and consecutive recruitment stages of the seedlings. We recorded the number of seeds deposited in seed traps to assess the local seed-deposition pattern and the abundance and distribution of seedlings and saplings to evaluate the spatial pattern of recruitment. More seeds were removed and deposited at the forest edge than in the interior. The number of deposited seeds decreased with distance from the fruiting tree and was spatially clustered in both habitat types. The density of 1-yr-old seedlings and saplings was higher at forest edges, whereas the density of 2-yr-old seedlings was similar in both habitat types. While seedlings were almost randomly distributed, seeds and saplings were spatially clustered in both habitat types. Our findings demonstrate systematic changes in spatial patterns of recruits across the plant regeneration cycle and suggest that the differential effects of biotic and abiotic factors determine plant recruitment at the edges and in the interior of tropical montane forests. These differences in the spatial distribution of individuals across recruitment stages may have strong effects on plant community dynamics and influence plant species coexistence in disturbed tropical forests.     

动物与植物种子更新的关系Ⅱ．动物对种子的捕食、扩散、贮藏及与幼苗建成的关系

植物的繁殖体总是面临来自各类生物（如昆虫、脊椎动物、真菌）的捕食风险。因动物捕食引起的种子死亡率影响植物的适合度、种群动态、群落结构和物种多样性的保持。种子被捕食的时间和强度成为植物生活史中发芽速度、地下种子库等特征的主要选择压力,而种子大小、生境类型等因素也影响动物对植物种子的捕食。捕食者饱和现象被认为是植物和种子捕食者之间的高度协同进化作用的结果,是限制动物破坏种子、提高被扩散种子存活率的一种选择压力。大部分群落中的大多数植物种子被动物扩散。种子扩散影响种子密度、种子被捕食率、病原体攻击率、种子与母树的距离、种子到达的生境类型以及建成的植株将与何种植物竞争,从而影响种子和幼苗的存活,最终影响母树及后代植物的适合度。种子被动物扩散后的分布一般遵循负指数分布曲线,大多数种子并没有扩散到离母树很远的地方。捕食风险、生境类型、植被盖度均影响动物对种子的扩散。植物结实的季节和果实损耗的过程也体现了其对扩散机会的适应。许多动物有贮藏植物种子的行为。动物贮藏植物繁殖体的行为,一方面调节食物的时空分布,提高了贮食动物在食物缺乏期的生存概率;另一方面也为种子萌发提供了适宜条件,促进了植物的扩散。于是,植物与贮食动物形成了一种协同进化关系,这种关系可能是自然界互惠关系（mutualism)的一种。影响幼苗存活和建成的因子包括种子贮蒇点的微生境、湿度、坡向、坡度、林冠盖度等。许多果食性动物吃掉果肉后,再将完好的种子反刍或排泄出来。种子经动物消化道处理后,发芽率常有所提高。     

动物与植物种子更新的关系I.对象,方法及意义

大多数植物靠种子库来更新。种子从离开母体到建成幼苗的过程中始终受动物活动的影响。动物对种子既有取食消耗的不利一面,又有将其扩散到适于发芽的安全地点的有利一面,二者处于一种利弊权衡状态。研究动物与植物种子更新的关系,有助于揭示种群动态的机制,理解动物和植物之间协同进化的规律,了解生态系统演替及其组分之间的关系,认识取食种子的动物在生态系统中的作用,进而为生态系统维持和生物多样性保护提供科学依据及有效     

Exploring the interaction of avian frugivory and plant spatial heterogeneity and its effect on seed dispersal kernels using a simulation model

Seed dispersal by avian frugivores is one of the key processes influencing plant spatial patterns, but may fail if there is disruption of plant–frugivore mutualisms, such as decline in abundance of dispersers, fragmentation of habitat, or isolation of individual trees. We used simulation model experiments to examine the interaction between frugivore density and behaviour and the spatial arrangement of fruiting plants and its effect on seed dispersal kernels. We focussed on two New Zealand canopy tree species that produce large fruits and are dispersed predominantly by one avian frugivore (Hemiphaga novaeseelandiae). Although the mean seed dispersal distance decreased when trees became more aggregated, there were more frugivore flights between tree clusters, consequently stretching the tails of the dispersal kernels. Conversely, when trees were less aggregated in the landscape, mean dispersal distances increased because seeds were deposited over larger areas, but the kernels had shorter tails. While there were no statistically meaningful changes in kernel parameters when frugivore density changed, decreases in density did cause a proportional reduction in the total number of dispersed seeds. However, birds were forced to move further when fruit availability and fruit ripening were low. Sensitivity analysis showed that dispersal kernels were primarily influenced by the model parameters relating to disperser behaviour, especially those determining attractiveness based on distance to candidate fruiting trees. Our results suggest that the spatial arrangement of plants plays an important role in seed dispersal processes – although tree aggregation curbed the mean seed dispersal distance, it was accompanied by occasional long distance events, and tree dispersion caused an increase in mean dispersal distance, both potentially increasing the probability of seeds finding suitable habitats for germination and growth. Even though low frugivore densities did not cause dispersal failure, there were negative effects on the quantity of seed dispersal because fewer seeds were dispersed.     

Changes of effective gene dispersal distances by pollen and seeds across successive life stages in a tropical tree

Pollen and seed dispersal are the two key processes in which plant genes move in space, mostly mediated by animal dispersal vectors in tropical forests. Due to the movement patterns of pollinators and seed dispersers and subsequent complex spatial patterns in the mortality of offspring, we have little knowledge of how pollinators and seed dispersers affect effective gene dispersal distances across successive recruitment stages. Using six highly polymorphic microsatellite loci and parentage analyses, we quantified pollen dispersal, seed dispersal, and effective paternal and maternal gene dispersal distances from pollen‐ and seed‐donors to offspring across four recruitment stages within a population of the monoecious tropical tree Prunus africana in western Kenya. In general, pollen‐dispersal and paternal gene dispersal distances were much longer than seed‐dispersal and maternal gene dispersal distances, with the long‐distance within‐population gene dispersal in P. africana being mostly mediated by pollinators. Seed dispersal, paternal and maternal gene dispersal distances increased significantly across recruitment stages, suggesting strong density‐ and distance‐dependent mortality near the parent trees. Pollen dispersal distances also varied significantly, but inconsistently across recruitment stages. The mean dispersal distance was initially much (23‐fold) farther for pollen than for seeds, yet the pollen‐to‐seed dispersal distance ratio diminished by an order of magnitude at later stages as maternal gene dispersal distances disproportionately increased. Our study elucidates the relative changes in the contribution of the two processes, pollen and seed dispersal, to effective gene dispersal across recruitment. Overall, complex sequential processes during recruitment contribute to the genetic make‐up of tree populations. This highlights the importance of a multistage perspective for a comprehensive understanding of the impact of animal‐mediated pollen and seed dispersal on small‐scale spatial genetic patterns of long‐lived tree species.     

Effects of habitat alteration on the effectiveness of plant-avian seed dispersal mutualisms: Consequences for plant regeneration

Anthropogenic habitat alteration may affect the dispersal service provided by avian seed dispersers, ultimately causing regeneration collapse, through a decay in both the quantitative (seed removal) and qualitative (seed arrival to safe sites) components of seed dispersal effectiveness. However, despite its implications for management in real-world landscapes, few studies have investigated the shifts in components of seed dispersal effectiveness resulting from habitat alteration. We advocate the use of stage-specific transition probabilities, combined with data on seed shadows and bird abundance and mobility, for a mechanistic inference of the consequences for recruitment of the disruption of plant-frugivore mutualism in altered habitats. Such an approach allows the identification of regeneration bottlenecks, evaluates the differential contribution to recruitment of quantitative and qualitative components of seed dispersal, and provides the means to compare seed dispersal limitation. We exemplify our conceptual approach with studies of seed dispersal and recruitment in the wild olive tree in unaltered and severely altered adjacent sites. We show that simplification of the habitat substantially affected bird abundance, diversity and mobility, which caused a reduction in fruit removal and a concomitant simplification of the seed shadows compared to the unaltered site. Linked to these shifts, postdispersal seed survival and seedling emergence and survival were affected. The final outcome of habitat alteration was the collapse of the regeneration dynamics with very few seeds escaping the influence of maternal plants and reaching the safest sites for recruitment (dispersal limitation). As predicted, the collapse in the regeneration dynamics resulted from severe decays in the quantitative but especially in the qualitative components of seed dispersal effectiveness. Management of fleshy-fruited plant populations in altered habitats should thus pay attention to landscape elements that promote frugivore abundance, diversity and mobility and that alleviate the dispersal limitation.     

How intraspecific variation in seed‐dispersing animals matters for plants

Seed dispersal by animals is a complex phenomenon, characterized by multiple mechanisms and variable outcomes. Most researchers approach this complexity by analysing context‐dependency in seed dispersal and investigating extrinsic factors that might influence interactions between plants and seed dispersers. Intrinsic traits of seed dispersers provide an alternative way of making sense of the enormous variation in seed fates. I review causes of intraspecific variability in frugivorous and granivorous animals, discuss their effects on seed dispersal, and outline likely consequences for plant populations and communities. Sources of individual variation in seed‐dispersing animals include sexual dimorphism, changes associated with growth and ageing, individual specialization, and animal personalities. Sexual dimorphism of seed‐dispersing animals influences seed fate through diverse mechanisms that range from effects caused by sex‐specific differences in body size, to influences of male versus female cognitive functions. These differences affect the type of seed treatment (e.g. dispersal versus predation), the number of dispersed seeds, distance of seed dispersal, and likelihood that seeds are left in favourable sites for seeds or seedlings. The best‐documented consequences of individual differences associated with growth and ageing involve quantity of dispersed seeds and the quality of seed treatment in the mouth and gut. Individual specialization on different resources affects the number of dispersed plant species, and therefore the connectivity and architecture of seed‐dispersal networks. Animal personalities might play an important role in shaping interactions between plants and dispersers of their seeds, yet their potential in this regard remains overlooked. In general, intraspecific variation in seed‐dispersing animals often influences plants through effects of these individual differences on the movement ecology of the dispersers. Two conditions are necessary for individual variation to exert a strong influence on seed dispersal. First, the individual differences in traits should translate into differences in crucial characteristics of seed dispersal. Second, individual variation is more likely to be important when the proportions of particular types of individuals fluctuate strongly in a population or vary across space; when proportions are static, it is less likely that intraspecific differences will be responsible for changes in the dynamics and outcomes of plant–animal interactions. In conclusion, focusing on variation among foraging animals rather than on species averages might bring new, mechanistic insights to the phenomenon of seed dispersal. While this shift in perspective is unlikely to replace the traditional approach (based on the assumption that all important variation occurs among species), it provides a complementary alternative to decipher the enormous variation observed in animal‐mediated seed dispersal.     

Frugivory and Seed Dispersal by Northern Pigtailed Macaques (Macaca leonina), in Thailand

Tropical rain forest conservation requires a good understanding of plant–animal interactions. Seed dispersal provides a means for plant seeds to escape competition and density-dependent seed predators and pathogens and to colonize new habitats. This makes the role and effectiveness of frugivorous species in the seed dispersal process an important topic. Northern pigtailed macaques (Macaca leonina) may be effective seed dispersers because they have a diverse diet and process seeds in several ways (swallowing, spitting out, or dropping them). To investigate the seed dispersal effectiveness of a habituated group of pigtailed macaques in Khao Yai National Park, Thailand, we examined seed dispersal quantity (number of fruit species eaten, proportion in the diet, number of feces containing seeds, and number of seeds processed) and quality (processing methods used, seed viability and germination success, habitat type and distance from parent tree for the deposited seeds, and dispersal patterns) via focal and scan sampling, seed collection, and germination tests. We found thousands of seeds per feces, including seeds up to 58 mm in length and from 88 fruit species. Importantly, the macaques dispersed seeds from primary to secondary forests, via swallowing, spitting, and dropping. Of 21 species, the effect of swallowing and spitting was positive for two species (i.e., processed seeds had a higher % germination and % viability than control seeds), neutral for 13 species (no difference in % germination or viability), and negative (processed seeds had lower % germination and viability) for five species. For the final species, the effect was neutral for spat-out seeds but negative for swallowed seeds. We conclude that macaques are effective seed dispersers in both quantitative and qualitative terms and that they are of potential importance for tropical rain forest regeneration.