脊椎动物传播植物肉质果中的次生物质及其生态作用

在种子植物-动物的互惠关系中,植物果实成熟后需要吸引种子传播者取食果实,传播其种子至适宜萌发的生境,同时又要防御种子捕食者过度消耗种子。果实内的次生物质(如:配糖生物碱、大黄素、辣椒素)在此过程中起到重要的调控作用。依赖脊椎动物传播的肉质果中往往含有与植物茎、叶内相同的次生物质,其种类繁多,主要分为含氮化合物、酚类化合物和萜类化合物。未成熟果实内富含次生物质(如:单宁、大黄素),主要保护未成熟种子不被潜在的捕食者和食果动物取食,这些次生物质的含量通常随果实成熟而降低;其它次生物质(如:脱辅基类胡萝卜素)的含量随果实成熟而增多,可能起到吸引食果动物的作用。在对脊椎动物捕食的抵御中,果实内不同类型的次生物质促使成熟果实对所有脊椎动物都有毒性(专毒性)或者仅对种子捕食者有毒性(泛毒性)。肉质果内的次生物质对植物-食果动物相互关系的调控作用,还可以通过调节动物取食频次和数量、抑制和促进种子萌发、改变种子在肠道的滞留时间、吸引传播者等生态作用而实现。某种次生物质往往集多种生态作用于一身。目前对肉质果内次生物质与脊椎动物相互关系的探讨还不够深入。未来研究需要综合考虑植物次生物质与果实生理生化、形态学等特征对食果者的综合调控机理;次生物质在种子传播后的调控作用对植物种群或群落结构和分布格局的影响;从动植物协同进化角度探讨植物次生物质的产生、防御和吸引策略与脊椎动物对果实的选择和消费之间的关系等。开展脊椎动物传播肉质果实中次生物质的研究,对完善种子传播机制、植物繁殖和更新格局,丰富动植物相互作用、协同进化理论具有重要的意义。     

种子捕食者对针叶树种子的捕食模式及二者的防御适应

在寒温带地区,针叶树种子因其油脂和蛋白含量高,成为许多动物的重要食物来源。在针叶林中,针叶树种子捕食者主要存在2种捕食方式,即传播前捕食和传播后捕食。通常,动物捕食种子的过程被认为是树种实现更新的重要途径。种子为躲避捕食者会形成相应的防御策略,种子的大小、营养、纤维含量、次级代谢产物等特征,通常也是植物种子防御捕食者捕食的策略。同样,这些防御策略一方面会影响种子捕食者的种群动态和取食行为,另一方面也可促使种子捕食者形成反对种子防御的适应对策。因此,研究种子捕食者捕食针叶树种子模式对于进一步认识针叶林中动物取食行为具有重要意义,同时,研究针叶树种子的防御及种子捕食者对其防御的适应策略,将有助于了解动植物之间相互作用。     

黄檗的更新特点及食果实鸟类对其种子的传播

在帽儿山实验林场和哈尔滨实验林场 ,黄檗母树下没有幼苗分布 ,不能进行天然更新 ,需要靠食果实鸟类等将果实和种子传播至远离母树的其他林下。捕食黄檗果实的食果实鸟类有 9种。其中 6种是食果肉鸟类 ,吞入果实后 ,消化果肉 ,而种子完整地随粪便排出而得以传播。其余 3种是食种子鸟类 ,没有传播作用。果实在鸟类消化道内的滞留时间达 2 0～ 30min ,具有很长的潜在传播距离。将鸟类消化后的种子与完整果实和人为去果肉种子进行萌发对比实验 ,消化后种子的累计萌发率与其余二者之间均没有显著性差异 ,说明食果实鸟类的消化 (道 )过程对种子萌发没有明显影响 ,同时证明果肉对种子萌发率没有显著影响 ,果肉中不含萌发抑制物质。黄檗提供多种鸟类以食物 ,而鸟类也同时以多种肉质果植物为食物。因此食果实鸟类和肉质果植物 (包括黄檗 )之间形成了松散的互利共生关系     

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。     

帽儿山地区秋季食果实鸟类的食性与迁徙

野外研究于2002和2003年秋季在帽儿山地区进行。帽儿山地区秋季可供鸟类取食的肉质果植物种类繁多,黄檗(Phellodendronamurense)和山荆子(Malusbaccata)是最主要的两种植物。帽儿山地区迁徙性食果实鸟类共9种,包括夏候鸟和旅鸟5种,冬候鸟4种。主要的食果实鸟类是斑鸫(Turdusnaumanni)、灰背鸫(T.hortulorum)和北朱雀(Carpodacusroseus),前两种为食果肉鸟类,后一种是食种子鸟类。斑鸫主要取食黄檗果实,灰背鸫主要取食山荆子果实,北朱雀主要取食黄檗种子。食果实鸟类在秋季也捕食一些动物性食物。3种主要食果实鸟类的迁徙日期不同。灰背鸫最先迁离,斑鸫稍晚,北朱雀最后到此越冬。果实丰歉年对鸟类的迁徙有一定的影响。     

食果动物与被取食植物的相互关系:协同进化?

在种子传播过程中动植物是否存在协同进化关系一直是争论的焦点。有观点认为,植物通过食果动物对其种子的传播可能获得逃避种子捕食者、占据新的生境斑块和基因流动等好处,而动物通过消化果肉获得营养和能量作为回报,动植物彼此相互作用,进而可能建立协同进化关系。动植物之间还可能发生在种、属或科水平以上的多物种的多配协同进化,或者通过关键种的协同进化来带动其他食果动物和植物相关性状的进化。“果肉防御假说”则认为果肉原本是保护种子的防御组织,后来才进化成为吸引食果动物以促进种子传播的物质。然而,食果动物和植物一对一的协同进化的例证并不多见;适合种子萌发和生长的环境在时空上难以确定;食果动物和植物的进化速度不一致;植物与种子传播者的选择压力存在着高度的不对称和不平衡,加上环境因素的重要影响,这种选择压力受到极大的限制而有可能变得不显著。种子传播中动植物在进化意义上的关系尚需进一步研究。未来研究应对食果动物和植物关系的复杂性和多样性有足够的认识。通过对系统发育中相联系的不同种的动植物关系的比较研究来揭示动植物关系对物种分化的影响,有可能为检验食果动物和植物之间的协同进化关系提供新的证据。食果动物传播种子对植物群落动态变化的影响、动植物关系和生物多样性保护等仍将是该领域研究的热点。     

应用结构方程模型解析影响黄连木果实产量 和种子命运的因素

果实(种子)产量和质量是影响植物种群更新的重要因素。为了探明影响黄连木果实产量和种子命运的因素以及这些影响因素之间的相互作用, 作者于2009年对河南省济源市45株黄连木(Pistacia chinensis)结果样树的植株特征、果实特征、果实产量和种子命运等进行了测定, 并用结构方程模型进行综合分析。结果表明: (1)黄连木果实产量与树高、树冠面积和果序大小等特征成正相关, 但与胸径、果实大小相关性不显著; (2)与捕食者饱和假说的预测不一致, 单株果实产量对黄连木广肩小蜂(Eurytoma plotnikovi)的种子捕食率(即虫蛀率)无显著直接负向效应; (3)树高和果实大小对虫蛀率为显著直接正向效应, 胸径对虫蛀率为显著直接负向效应, 显示黄连木广肩小蜂对植株特征和果实特征有一定的选择能力; (4)空壳率与虫蛀率成显著负相关, 空壳果实越多, 越易逃避黄连木广肩小蜂的寄生, 空壳果实的存在对完好种子起到了一定保护作用, 可能是黄连木防御昆虫寄生的重要机制; (5)空壳率和虫蛀率对种子完好率有显著直接负向效应, 而胸径、果序大小和果实产量对完好率为间接正向效应, 树高和果实大小为间接负向效应。可见, 黄连木植株特征和果实特征均不同程度地影响其果实产量和昆虫寄生, 从而影响黄连木的种子质量和种群更新。     

果实生态学的概念、研究现状及研究方向

总结了果实生态学的概念及主要研究内容,对国内外的果实生态学研究领域的进展进行了综述,分析了这些研究尚存在的不足之处,并指出了未来研究的科学问题.迄今为止,果实生态学在果实与种子的关系、果实颜色与环境、果实化学成分与环境、群落果实构成式样的地理分布特征、果实及食果动物的协同进化等方面已经取得了若干进展.果实重量谱等物理性状的空间分异格局及机理、果实化学成分的时空变异格局、果实类型的组成式样及大尺度地理分布格局及其机制的探讨、果实传播与食果动物的协同进化等是未来亟待开展的工作.     

黑尾蜡嘴雀冬季对树木果实的取食作用

2009年11月至2011年2月,在江苏省南京市观察到黑尾蜡嘴雀冬季取食12种树木的果实及种子。该鸟通常咬破果皮或种皮,取食种子的胚和胚乳等营养物质,为种子捕食者。由于其他一些食果鸟类也同期取食有关树木的果实,并传播其种子,黑尾蜡嘴雀的取食对这些树木的种子传播的影响是有限的。在初冬季节,黑尾蜡嘴雀常在树冠层取食果实,而在深冬季节,则主要在地表取食落果。黑尾蜡嘴雀能够咬碎枫杨等坚果的坚硬果皮,可为一起在地表觅食的麻雀取食种子碎屑物提供便利条件。黑尾蜡嘴雀与麻雀之间的偏利取食关系属于首次报道。文中对于城市环境管理中合理利用树木落果,为黑尾蜡嘴雀等鸟类提供越冬食物等方面提出了相关建议。     

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。     

Effects of seed size and frugivory degree on dispersal by Neotropical frugivores

Large vertebrates are important elements of mutualistic interactions and provide positive impacts on plant population and community dynamics. Despite the increasing interest on vertebrate frugivory we are still not able to disentangle the real contribution of seed dispersal to Neotropical forest functioning. Consuming fruits does not imply effective seed dispersal and many variables, such as seed size and animal diet, may influence the outcome of plant-animal interactions. Here, we performed a comprehensive literature search on seed dispersal by Neotropical vertebrates (with a focus on primates) to closely approach their role as seed dispersers, hypothesizing frugivory degree and seed size as main drivers of fruit handling behavior and diversity of dispersed seeds. We found that the great majority of seeds manipulated by Neotropical primates, with exception to the seed predators pitheciins, were swallowed and passed intact through their gut. Larger seeds (>12 mm) tended of being ingested exclusively by primates and other large vertebrates, such as tapirs and peccaries. Furthermore, primate feeding guild had a great influence on the richness and sizes of seeds dispersed, as primarily frugivores dispersed more species and had higher probabilities of ingesting larger seeds when compared to other feeding guilds. Organizing available knowledge and filling the main knowledge gaps allowed us to validate common sense assumptions and ultimately draw new conclusions about the role played by primates together with other major frugivores in Neotropical forests.     

Sowing forests: a synthesis of seed dispersal and predation by agoutis and their influence on plant communities

Granivorous rodents have been traditionally regarded as antagonistic seed predators. Agoutis (Dasyprocta spp.), however, have also been recognized as mutualistic dispersers of plants because of their role as scatter-hoarders of seeds, especially for large-seeded species. A closer look shows that such definitions are too simplistic for these Neotropical animals because agoutis can influence plant communities not only through seed dispersal of large seeds but also through predation of small seeds and seedlings, evidencing their dual role. Herein, we summarize the literature on plant–agouti interactions, decompose agouti seed dispersal into its quantitative and qualitative components, and discuss how environmental factors and plant traits determine whether these interactions result in mutualisms or antagonisms. We also look at the role of agoutis in a community context, assessing their effectiveness as substitutes for extinct megafaunal frugivores and comparing their ecological functions to those of other extant dispersers of large seeds. We also discuss how our conclusions can be extended to the single other genus in the Dasyproctidae family (Myoprocta). Finally, we examine agoutis’ contribution to carbon stocks and summarize current conservation threats and efforts. We recorded 164 interactions between agoutis and plants, which were widespread across the plant phylogeny, confirming that agoutis are generalist frugivores. Seed mass was a main factor determining seed hoarding probability of plant species and agoutis were found to disperse larger seeds than other large-bodied frugivores. Agoutis positively contributed to carbon storage by preying upon seeds of plants with lower carbon biomass and by dispersing species with higher biomass. This synthesis of plant–agouti interactions shows that ecological services provided by agoutis to plant populations and communities go beyond seed dispersal and predation, and we identify still unanswered questions. We hope to emphasise the importance of agoutis in Neotropical forests.     

Importance of animal and plant traits for fruit removal and seedling recruitment in a tropical forest

The traits of animals and plants influence their interaction networks, but the significance of species' traits for the resulting ecosystem functions is poorly understood. A crucial ecosystem function in the tropics is seed dispersal by animals. While the importance of species' traits for structuring plant–frugivore networks is supported by a number of studies, no study has so far identified the functional traits determining the subsequent processes of fruit removal and seedling recruitment. Here, we conducted a comprehensive field study on fruit removal by frugivorous birds and seedling recruitment along an elevational gradient in the Colombian Andes. We measured morphological traits of birds (body mass, bill width, Kipp's index) and plants (plant height, crop mass, fruit width and seed mass) which we expected to be related to fruit removal and seedling recruitment. We tested 1) which bird and plant traits influence fruit removal, and 2) whether network metrics at plant species level, functional identities of frugivores (community‐based mean trait values) and/or plant traits were the main determinants of seedling recruitment. We found that large‐bodied bird species contributed more to fruit removal than small‐bodied bird species and that small‐sized fruits were more frequently removed than large‐sized fruits. Small plant species and plants with heavy seeds recruited more seedlings than did large plants and plants with light seeds. Network metrics and functional identities of seed dispersers were unrelated to seedling recruitment. Our findings have two important implications. First, large birds are functionally more important than small birds in tropical seed‐removal networks. Second, the detected tradeoff between fruit size and seed mass in subsequent recruitment processes suggests that the adaptability of forest plant communities to a loss of large frugivores is limited by life‐history constraints. Hence, the protection of large‐bodied frugivores is of primary importance for the maintenance of diverse tropical plant communities.     

Influence of type of avian frugivory on the fitness ofPistacia terebinthus L.

Summary The fruits ofPistacia terebinthus, a circum-Mediterranean tree/shrub, are consumed by an array of bird species that differ in feeding methods and in relative frequencies of visits to plants. In this study I document interindividual variation in the proportion of fruits consumed by three types of frugivores: legitimate dispersers, pulp-consumers and seed predators. The results show that the relative frequencies of each kind of frugivore notably influence the final reproductive output (absolute number of viable seeds dispersed) and in fact prevail over the effects of pre-dispersal factors acting on plant fitness. Those relative frequencies are not associated with any of the plant traits related to fitness, such as fruit crop size and the number of viable seeds produced, suggesting that the type of avian frugivory exerts a negligible, if not null, selective pressure on such plant attributes. Plant specialization to attract the most effective seed dispersers seems to be precluded, given the small scale at which the high variation in seed dispersal success takes place.     

Interactions between mammals and trees: predation on mammal-dispersed seeds and the effect of ambient food

The Janzen-Connell escape hypothesis predicts that the success of tree propagules increases with distance from the parent tree. Fleshy fruits that are transported in the guts of frugivores are believed to have evolved to facilitate the wide dispersal of seeds. However, some frugivores deposit seeds in latrines, thus creating aggregations of seeds that are large enough to attract seed predators and negate the advantages of dispersal. Raccoons (Procyon lotor) often produce large seed deposits since they habitually defecate in latrines. The survival of wild black cherry (Prunus serotina) seeds in simulated raccoon latrines was monitored in areas with natural levels of food availability and in areas to which supplemental food had been supplied to the primary seed predators. Dispersal of seeds by raccoons did not necessarily provide effective protection from post-dispersal seed predation at natural food levels. Once the resident seed predators had located the latrines, the majority of the seeds were quickly removed. However, seed removal from raccoon latrines was reduced significantly and dramatically by the addition of alternative food. This implies that raccoon latrines may represent safe sites for tree recruitment during periods of high food availability such as during masting events, thus providing conditional support for the escape hypothesis.     

Unexpected relationships and valuable mistakes: non‐myrmecochorous Prosopis dispersed by messy leafcutting ants in harvesting their seeds

Abstract Ants generally disperse seeds while feeding on fruits or structures attached to the seed. Seed dispersal as a by‐product of seed predation (dyszoochory) was recognized in specialized harvester ants, but not in ants predating seeds opportunistically. Leafcutting ants are the main herbivores in much of the Neotropics, and they have been reported to remove fruits and seeds, but their role as seed predators and dispersers has not been acknowledged. Prosopis flexuosa D.C. (Fabaceae, Mimosoideae) is the most abundant tree species in the central Monte Desert, Argentina, and it is likely to depend on secondary animal dispersal. Mammalian frugivores are usually considered its main dispersers, but the opportunity for dispersal may be small since the removal of fruits and seeds by seed predators is very intense. The objective of this study was to identify which ant species interact with P. flexuosa fruits and to evaluate their relative importance as seed predators and dispersers. In a field experiment, whole and segmented pods were offered and several ant species exploiting the fruits were identified. Additionally, all pod segments remaining around nests of the three ant species able to remove them (the leafcutters Acromyrmex lobicornis Emery and Acromyrmex striatus Roger, and Pheidole bergi Mayr) were examined during and after the P. flexuosa primary dispersal season. Up to 753 pod segments and 90 sound seeds were found accumulated in a circle of 1 m radius over nests of A. lobicornis, and even more in an examined trail. Acromyrmex striatus left a smaller proportion of sound seeds and P. bergi left a smaller number of pod segments. All tendencies were similar during shorter known periods of accumulation. Leafcutting ants are acting as important seed predators, and ‘by mistake’ may be dispersing a key non‐myrmecochorous tree. This is an unexplored path in the seed dispersal cycle of P. flexuosa that challenges the tendency to predict interactions based on classifications made with other goals.     

The Role of Arboreal Seed Dispersal Groups on the Seed Rain of a Lowland Tropical Forest1

Most tropical plants produce fleshy fruits that are dispersed primarily by vertebrate frugivores. Behavioral disparities among vertebrate seed dispersers could influence patterns of seed distribution and thus forest structure. This study investigated the relative importance of arboreal seed dispersers and seed predators on the initial stage of forest organization–seed deposition. We asked the following questions: (1) To what degree do arboreal seed dispersers influence the species richness and abundance of the seed rain? and (2) Based on the plant species and strata of the forest for which they provide dispersal services, do arboreal seed dispersers represent similar or distinct functional groups? To answer these questions, seed rain was sampled for 12 months in the Dja Reserve, Cameroon. Seed traps representing five percent of the crown area were erected below the canopies of 90 trees belonging to nine focal tree species: 3 dispersed by monkeys, 3 dispersed by large frugivorous birds, and 3 wind‐dispersed species. Seeds disseminated by arboreal seed dispersers accounted for ca 12 percent of the seeds and 68 percent of the seed species identified in seed traps. Monkeys dispersed more than twice the number of seed species than large frugivorous birds, but birds dispersed more individual seeds. We identified two distinct functional dispersal groups, one composed of large frugivorous birds and one composed of monkeys, drop dispersers, and seed predators. These groups dispersed plants found in different canopy strata and exhibited low overlap in the seed species they disseminated. We conclude it is unlikely that seed dispersal services provided by monkeys could be compensated for by frugivorous birds in the event of their extirpation from Afrotropical forests.     

Ecology of the fruit-colour polymorphism in Rubus spectabilis

Although some studies have focused on the colour polymorphisms of flowers and fruits, little is known of their ecological and evolutionary significance. We investigated the potential contribution of several factors to the maintenance of fruit-colour polymorphism in Rubus spectabilis, a common shrub in the temperate rainforests of southeast Alaska. Fruits occur in two colours (red and orange), whose frequencies vary geographically. The two colour morphs have similar size, weight, seed load and nutrient composition. Colour preferences of avian frugivores, in the aviary and in the field, varied among individuals, but the majority favoured red fruits. Seed predators (mostly rodents) did not discriminate between seeds from different morphs. The effect of seed passage through the digestive tract of frugivores (birds and bears) on germination was similar for both morphs, although there were significant differences among frugivores. The type of soil on which the seeds are deposited influenced their germination behaviour, suggesting that some soils could favour one morph over the other. Such differences may contribute to the regional differences in frequencies of the two morphs. This study emphasizes the need to investigate fruit and seed characteristics that correlate with fruit colour; the colour preferences of consumers is only one of several selection pressures that determine the frequency distribution of fruit colours. This revised version was published online in July 2006 with corrections to the Cover Date.