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。     

凋落物和土壤覆盖对动物取食和搬运辽东栎种子的影响

在六盘山区的华北落叶松人工林,研究了清除凋落物、凋落物覆盖和土壤覆盖(以不清除凋落物直接将种子投放于森林地表为对照)等处理对动物取食和搬运辽东栎种子的影响.结果表明:种子释放3d后,凋落物和土壤覆盖处理种子均具有较高的留存率(分别为10.7％和7.0％);释放14 d后,土壤覆盖处理种子的留存率仍最高(0.7％),但凋落物覆盖处理种子的留存率为0.凋落物和土壤覆盖处理种子的就地取食率很高(分别为45.9％和41.5％);清除凋落物处理种子的就地取食率最低(27.0％),但其被搬运后的取食率最高(49.8％);凋落物覆盖、清除凋落物和土壤覆盖处理种子被动物搬运后的埋藏率均显著高于对照(P＜0.01).种子被动物搬运后集中分布于5 m以内,尤其在＜1 m和l～2m两个距离组的分布频率更高;种子被搬运后取食的平均距离大干埋藏的平均距离,以土壤覆盖和凋落物覆盖处理最大,分别为2.38 m±0.55 m和1.44m±0.26 m.     

啮齿动物对不同林木种子的搬运和取食微生境选择机制

不同植物种子依靠不同的方式实现扩散,啮齿动物对林木种子搬运后在取食点微生境和贮藏方式的选择存在偏好,研究其贮藏行为与微生境的关系是探究幼苗建成的关键。在秦岭中段火地塘林区,采用标签标记法,以锐齿槲栎、华山松和油松种子为材料,探究了小型啮齿动物对松栎混交林建群种种子扩散过程的影响。结果表明:1)油松种子原地取食率显著高于锐齿槲栎和华山松种子,且啮齿动物更倾向于搬运后取食(60%)和埋藏(4.33%)华山松种子,搬运后取食距离也为华山松最大(2.49 m);锐齿槲栎小种子被搬运后埋藏的距离最大(4.92 m)。2)除华山松种子外,其他类型种子被搬运后单个取食的比例均在85%以上;油松种子不存在埋藏点,而其他类型种子90%以上均以单个形式被埋藏。3)大部分种子被啮齿动物搬运后选择在裸地丢弃;锐齿槲栎大种子(87.5%)、小种子(78.57%)和华山松种子(53.33%)较大比例被啮齿动物埋藏在灌丛下方,埋藏在裸地的种子较少。4)大部分种子在灌丛下方被取食,仅华山松种子被啮齿动物搬运到洞穴取食;除油松种子被大量原地取食外,其他类型种子被搬运到取食点的种子比例基本呈现由微生境植被复杂到简单(灌丛—草丛—灌丛边缘—裸地)而逐渐减小的趋势。种子的营养价值及取食和搬运过程中啮齿动物付出的成本是影响种子命运的关键性因子,且啮齿动物对种子埋藏和取食地点的微生境存在较明显的选择性。     

濒危植物银杉的结实特性及动物对果实的危害性

银杉结实量的年际波动很大,具有明显的大小年之分,一般在２～３个小年之后为１～２个大年。银杉母株的结实量在不同的母树间差别很大,平均单株的球果数仅为６６个。球果结实量在树冠南北侧有差异,南侧球果数占球果总数的６０％;球果结实量的垂直分异明显,树冠中、上部的球果数占总量的９０％以上。松鼠对树上尚未完全成熟的银杉果实的取食危害是长期性的,对银杉种子种群造成严重破坏。以１９９５年为例,被啃食的球果数占球果总数的比重可达１５％左右。试验表明,土壤种子库中地表的银杉种子几乎全部被动物取食,种子埋藏深度增加有利于逃避动物的取食。     

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。     

动物对花楸树种实的取食与传播

花楸树是我国东北林区重要的非木质资源树种,其种实既有自然散布方式,也有动物散布方式.本文通过对花楸树种实散布过程中动物活动特点的研究,探索动物取食和传播花楸树种实的规律及其对花楸树天然更新的影响.在2008和2009年花楸树果实成熟期,通过定期观察取食花楸树果实的鸟类及其取食方式,确定对花楸树果实有拜访行为的鸟类有8种,其中食果肉鸟类斑鸫、灰喜鹊和白背啄木鸟对花楸树种实有传播作用,它们对花楸树果实的拜访频率分别为54%、12%和7%,取食后第一落点集中于距离母树5～10 m之间（占68.2%）,其次为距离母树5 m以内（占27.3%）,距离母树10 m以外的比例很小（占4.5%）.果实在鸟类消化道内的滞留时间可达20 min,表明潜在传播距离会很长.人工摆放果实和种子试验表明,在不同生境地面摆放的果实6～7 d内全部消失,果实的取食者主要是啮齿类和地面取食的鸟类,取食率均较高（50%～70%）;种子的取食者为啮齿类、地面取食的鸟类和蚁类,取食率均较低（1%～5%）.花楸树为多种动物提供食物,而动物为花楸树传播种子,动物的取食对花楸树的天然更新有重要影响.     

原始老龄林内兴安落叶松种子命运的试验研究

兴安落叶松土壤种子库的支出主要有３个方面：种子萌发、动物取食和生活力丧失。兴安落叶松种子在６月份迅速萌发,占全年总萌发量的９０％。７月中下旬,土壤已经不存在有生活力的种子。在生长季初期,鼠类对种子取食率仅１％,到生长季中后期接近２０％。然而,此时幼苗发生基本结束,被食种子大部分为生活力很弱或丧失生活力者。红背Ｐｉｎｇ是主要的食种子者。落叶松在时间上逃避了鼠类取食种子。一般,鼠类仅就地取食１０％的落     

土壤埋藏深度和埋藏时间对辽东栎种子萌发的影响

以采自宁夏六盘山区的辽东栎(Quercus wutaishanica)种子为材料,研究了不同土壤埋藏深度(0、30和50 cm)和埋藏时间(2、4、6、8、10和12周)对种子萌发的影响,以探索辽东栎种子的安全贮藏条件和长期贮藏潜力,为辽东栎的种苗繁育和退化辽东栎灌丛及次生林的恢复实践提供参考.结果表明,尽管土壤埋藏处理前期(2周)种子萌发受到抑制,但随着埋藏时间的延长,萌发率提高,萌发速率加快,且萌发指数和活力指数均呈波动性增大趋势,埋藏12周后,30和50 cm埋深处理种子的萌发率分别为90.1％和96.8％.种子霉烂率表现为随埋藏深度的增加而提高的趋势,且各埋深处理种子的霉烂率随埋藏时间的延长而提高.种子埋藏2周后,开始出现自动萌发现象;埋藏12周后,30和50 cm埋深处理种子的自动萌发率分别为24.5％和43.0％.     

广西扶绥黑叶猴的主要食源植物及其粗蛋白含量

食物的营养物质是影响动物食物选择的生态因子。研究动物食物的营养组成及其对野生动物取食的影响对野生动物的保护具有重要的参考意义。为了研究喀斯特环境中食物粗蛋白对黑叶猴食物选择的影响,2006年1月-12月在广西扶绥自然保护区, 采用焦点动物取样法和连续记录法研究了片断化栖息地中黑叶猴的觅食行为,同时采集黑叶猴取食部位样本共40种190份,用凯氏定氮法测量其粗蛋白含量。结果表明,黑叶猴取食的食物包括嫩叶、成熟叶、花、果和种子等部位;食源植物的生活型包括了乔木、灌木、藤本和草本。食物的平均粗蛋白含量为12.7%,有明显的季节性变化。4月份样品的月平均粗蛋白含量全年最高,高达19.1%,而最低在6月份,仅为5.4%。食物的粗蛋白含量在前半年和后半年之间没有显著的差异（z= -1.28,p= 0.20）,在雨季和旱季之间亦无差异（z= -1.22,p= 0.22）;不同食源植物部位的平均粗蛋白含量不同,嫩叶的年平均粗蛋白含量最高,为13.7%,其次是成熟叶和花,分别是13.0%和12.1%,果实最低,仅为9.6%;不同生活型的食源植物粗蛋白含量不同,藤本的粗蛋白含量最高,为 14.1%,草本的最低,仅为10.1%,乔木和灌木分别是11.3%。不同食源植物的取食时间与其粗蛋白含量没有显著的相关关系（p>0.05）,粗蛋白含量不是影响黑叶猴取食的主要因素。     

食种子动物对三种锦鸡儿属植物繁殖更新的影响

锦鸡儿属(Caragana)植物是黄土高原植被的重要组分,具有固氮、耐旱的特性,对黄土丘陵沟壑区的植被恢复具有重要意义。作者于2002–2003年对该属本地种白毛锦鸡儿(C.licentiana)和甘蒙锦鸡儿(C.opulens)及外来种中间锦鸡儿(C.intermedia)种子被动物取食情况进行了比较研究。2003年白毛锦鸡儿种子散落前的虫害率为88.2%,甘蒙锦鸡儿为29.7%±1.7%,中间锦鸡儿为43.2%±4.8%。种子落地后,白毛锦鸡儿和甘蒙锦鸡儿的种子被小啮齿动物取食的比例在各种微生境下均为100%,且取食速度很快,在种子放置后4d内即全部被取食。中间锦鸡儿种子被取食率则因微生境的不同而不同:在灌丛下被取食率为100%,在半阳坡的开阔草地上为85.0%±10.0%,在种植植物未成功的水平阶地上为31.0%±8.7%,且取食速度相对较慢。研究结果表明,食种子动物的取食大大减少了白毛锦鸡儿和甘蒙锦鸡儿的种子量以及种子传播萌发的机会。但食种子动物可能充当了白毛锦鸡儿4.0%硬实种子的传播者,从而有助于白毛锦鸡儿实现种子途径的更新;中间锦鸡儿种子无论在散落前还是散落后受食种子动物的影响均很小。     

Effects of dung and seed size on secondary dispersal,seed predation,and seedling establishment of rain forest trees

Seeds dispersed by tropical, arboreal mammals are usually deposited singly and without dung or in clumps of fecal material. After dispersal through defecation by mammals, most seeds are secondarily dispersed by dung beetles or consumed by rodents. These post-dispersal, plant-animal interactions are likely to interact themselves, as seeds buried by dung beetles are less likely to be found by rodents than unburied seeds. In a series of three experiments with seeds of 15 species in central Amazonia (Brazil), we determined (1) how presence and amount of dung associated with seeds influences long-term seed fate and seedling establishment, (2) how deeply dung beetles bury seeds and how burial depth affects seedling establishment, and (3) how seed size affects the interaction between seeds, dung beetles, and rodents. Our overall goal was to understand how post-dispersal plant-animal interactions determine the link between primary seed dispersal and seedling establishment. On average, 43% of seeds surrounded by dung were buried by dung beetles, compared to 0% of seeds not surrounded by dung (n=2,156). Seeds in dung, however, tended to be more prone than bare seeds to predation by rodents. Of seeds in dung, probability of burial was negatively related to seed size and positively related to amount of dung. Burial of seeds decreased the probability of seed predation by rodents three-fold, and increased the probability of seedling establishment two-fold. Mean burial depth was 4 cm (0.5–20 cm) and was not related to seed size, contrary to previous studies. Probability of seedling establishment was negatively correlated with burial depth and not related to seed size at 5 or 10 cm depths. These results illustrate a complex web of interactions among dung beetles, rodents, and dispersed seeds. These interactions affect the probability of seedling establishment and are themselves strongly tied to how seeds are deposited by primary dispersers. More generally, our results emphasize the importance of looking beyond a single type of plant-animal interaction (e.g., seed dispersal or seed predation) to incorporate potential effects of interacting interactions.     

Predation and dispersal of large and small seeds of a tropical palm

Seed size may vary greatly among individuals within plant species. What effects the extremes of this variation have for seeds taken by small mammals are poorly understood. Not all seeds removed by small mammals are necessarily eaten. Small rodents are common seed predators, but they may disperse a significant proportion of seeds by scatter hoarding them via burial. Size-dependent predation and dispersal of seeds has not been directly tested within a plant species for tropical rodents. This study tested whether or not large and small nuts of Astrocaryum mexicanum (Palmae) differed in their fates due to handling by the spiny pocket mouse Heteromys desmarestianus (Heteromyidae). Exclosures were used to give small rodents exclusive access to A. mexicanum nuts. H. desmarestianus preferentially consumed large over small A. mexicanum nuts, but cached (in burrows and by scatter hoarding) similar proportions of these nuts by size. Small nuts tended to be buried farther away from exclosures than large nuts. Although sample sizes of buried nuts were small, the rodents retrieved all buried large nuts, but 30% of the small nuts remained buried long enough to germinate. I also examined predispersal predation by insects and found that insects appear to have no size preference for A. mexicanum nuts, but insect predation appears to hinder nut development. Thus, nuts attacked by insects develop to be significantly smaller, with a low proportion of undamaged endosperm, than uninfested nuts. It is hypothesized that the preferential predation of large A. mexicanum nuts by H. desmarestianus is a response by these rodents to insect predation.     

Post-dispersal seed predation: consequences for plant demography and evolution

Post-dispersal seed predation is only one of many factors underlying plant demography and evolution. Nevertheless, the generalist feeding habits of many post-dispersal seed predators and the limited ability of plants either to compensate for or to respond to post-dispersal seed losses directly suggest that post-dispersal seed predation may have a considerable impact on plant populations. Seed predators probably have little direct influence on the demography of plants that regenerate exclusively by vegetative means or are buffered by a large active seed bank, but such species are only a minority in most plant communities.In general, ants are significant post-dispersal seed predators in arid and semi-arid ecosystems while they act mainly as seed dispersers rather than as predators in temperate ecosystems. Although studies have probably underestimated the importance of invertebrates and birds as seed predators, rodents appear to have greater potential to influence seed dynamics, and are particularly important in temperate ecosystems. For example, production of mast seed crops is more effective at satiating specialist invertebrate seed predators than generalist vertebrates, and recruitment may be limited by post-dispersal seed predation even during mast years.Both spatial variation in post-dispersal seed predation and differences in predation between species are important elements which facilitate the coexistence of different plant species. Where microsites are limiting, selective post-dispersal seed predators can influence pre-emptive competition for these microsites. Seed size determines the extent of density-dependent predation and the exploitation of buried seed. This suggests that post-dispersal seed predators may also play a role in the evolution of seed characteristics. However, conclusions regarding the ecological and evolutionary impact of post-dispersal seed predators will remain speculative without a more substantial empirical base.     

Variability in post-dispersal seed predation in deciduous woodland: relative importance of location,seed species,burial and density

The considerable variability found in post-dispersal seed predation and the absence of consistent directional trends (e.g., with reference to seed size) has made it difficult to predict accurately the impact of seed predators on plant communities. We examined the variation attributable to location, seed density and seed burial on the removal of seeds of three tree species: Fraxinus excelsior, Taxus baccata and Ulmus glabra. Experiments were undertaken in five deciduous woodlands in Durham, U.K., and the relative importance of vertebrate and invertebrate seed predators was assessed using selective exclosures. In all five woodlands, seed removal was greatest from treatments to which vertebrates had access, and losses attributable to invertebrates were negligible. Rodents, in particular Apodemus sylvaticus (Muridae) and Clethrionomys glareolus (Cricetidae), were the principal seed consumers in these woodlands. Unidentified vertebrate seed predators (probably birds, rabbits and/or squirrels) appeared to be significant seed removers in three of the five woodlands. Rates of removal differed among the three tree species, increasing in the following order Fraxinus < Taxus < Ulmus but were not related to seed mass. The major effect influencing rates of seed removal was seed burial, which halved rates of seed removal overall. The effect of seed burial was a function of seed size. The larger seeds of Taxus realising little benefit from seed burial whereas encounter of the smaller Ulmus seeds fell by almost two-thirds. Removal was density-dependent for all three species. However, the relative increase in seed encounter through an increase in seed density was a negative function of seed size. This suggests that, for large seeds, the opportunity to escape seed predation via burial or reduced seed density is limited. These results reveal a number of parallels with other studies of post-dispersal predation and identify several generalities regarding the interaction between plants and post-dispersal seed predators. Comparison of the seed predation results with actual seedling distributions suggests that seed predators may influence regeneration of Ulmus glabra but probably play a lesser role in the dynamics of Taxus baccata and Fraxinus excelsior.     

Seed Dispersal by Monkeys and the Fate of Dispersed Seeds in a Peruvian Rain Forest1

Primary seed dispersal by two species of monkeys and the effects of rodents and dung beetles on the fate of dispersed seeds are described for a rain forest in southeastern Perú. During the six-month study period (June–November 1992) spider monkeys (Ateles paniscus) dispersed the seeds of 71 plant species, whereas howler monkeys (Alouatta seniculus) dispersed seeds of 14 species. Spider and howler monkeys also differed greatly in their ranging behavior and defecation patterns, and as a consequence, produced different seed rain patterns. Monkey defecations were visited by 27 species of dung beetles (Scarabaeidae). Dung beetles buried 41 percent of the seeds in the dung, but the number of seeds buried varied greatly, according to seed size. Removal rates of unburied seeds by rodents varied between 63–97 percent after 30 d for 8 plant species. The presence of fecal material increased the percentage of seeds removed by seed predators, but this effect became insignificant with time. Although seed predators found some seeds buried in dung balls (mimicking burial by dung beetles), depth of burial significantly affected the fate of these seeds. Less than 35 percent of Brosimum lactescens seeds buried inside dung balls at a depth of 1 cm remained undiscovered by rodents, whereas at least 75 percent of the seeds escaped rodent detection at a depth of 3 cm and 96 percent escaped at 5 cm. Both dung beetles and rodents greatly affected the fate of seeds dispersed by monkeys. It is thus important to consider postdispersal factors affecting the fate of seeds when assessing the effectiveness of frugivores as seed dispersers.     

Early reduction of post-fire recruitment of Pinus nigra by post-dispersal seed predation in different time-since-fire habitats

This study analyses the effects of post-dispersal predation of Pinus nigra seeds on the initial recruitment of this species in areas burned by large wildfires, where P. nigra shows very low regeneration. In three different habitats obtained in a gradient of time since fire in Catalonia (NE Spain), we have evaluated the effects of seed predators (ants, rodents and birds) on post-dispersal seed removal and early seedling establishment of P. nigra by using selective exclosures limiting their access to seeds. Ants were the most efficient seed predator group, followed by rodents and birds. The contribution of each group to overall predation showed large seasonal variations. The first seeds dispersed in winter were mainly predated by rodents, which also registered their highest abundance in this season of the year. In spring, at the end of the natural dissemination period of P. nigra seeds, ants became the major predators, this fact coinciding with their increased abundance. Birds showed the lowest predation values. In the seedling establishment experiment, only in the exclusion treatment of the three predator groups was there initial establishment in all habitats, especially in the recently burned area, where there was seedling establishment in all exclusion treatments. The post-dispersal seed predation by different animal groups and low seedling emergence in the different habitats obtained in this study, together with the low seed availability of P. nigra seeds in burned areas, do not predict a favourable outlook for the natural post-fire recolonization of this species, which might even affect its overall distribution area in the region.     

亚热带林区啮齿动物对樱桃种子捕食和搬运的作用格局

在都江堰林区,通过在原生林、次生林和灌丛3类生境释放野生和栽培樱桃(Cerasus pseudocerasus)的种子,研究了啮齿动物对樱桃种子的捕食及其对樱桃种群更新的作用。结果表明,啮齿动物对樱桃两类种子的搬运无显著差异,而啮齿动物对野生樱桃种子的收获则明显快于栽培樱桃种子,且在3类生境均有类似的趋势。这说明啮齿动物偏爱于收获具有较高收益(种仁重/种子重)的野生樱桃种子。啮齿动物在小于10 d的时间尺度收获了所有释放的樱桃种子,其中,70%以上为啮齿动物所搬运。春季食物的匮乏可能是导致啮齿动物对樱桃种子有较大捕食压力的主要原因,而生境类型间的差异较小。因此,啮齿动物是都江堰林区樱桃地表种子的主要捕食者,它们对野生樱桃种子的选择性捕食和搬运能影响樱桃种子/果实的进化及其种群更新。     

Seed Dynamics of Early and Late Successional Tree Species in Tropical Abandoned Pastures: Seed Burial as a Way of Evading Predation

We explored different treatments to enhance the probability of sowed seeds of two early successional (ES, Cecropia obtusifolia and Ochroma pyramidale) and two late successional (LS, Brosimum costaricanum and Dialium guianense) species to escape predation and germinate in abandoned cattle‐raising pasture fields in Southeastern Mexico. ES species were sown in groups of 50 seeds under three treatments: invertebrate exclusion, burial, and exposition to seedeaters. LS species were sown in groups of 10 seeds under three treatments: vertebrate exclusion, burial, and exposition to seedeaters. We registered seed predation and germination 2, 4, 8, 16, 32, and 64 days after the initial sowing. Overall, ES showed higher predation rates (mean ± SE = 0.45 ± 0.07 seed seed^?1 day^?1; n = 3) than LS species (0.09 ± 0.02 seed seed^?1 day^?1). Cecropia obtusifolia was completely predated in all treatments after 8 days. Burial and exclusion treatments reduced final predation in circa 6% for O. pyramidale, relative to that of exposed seeds (85% after 8 days); most germination occurred in buried seeds (3.7%). In B. costaricanum, burial enabled germination by 10%; exposed and excluded seeds were removed 100%. Dialium guianense showed 12% germination in buried seeds and circa 20% of the seeds were not removed after 64 days. Direct sowing would be a recommended rainforest restoration practice for species with relatively large seeds if deposited in groups and buried. Studies which address variation across numerous sites are necessary in order to generate more consistent seed predation patterns and rainforest restoration principles in tropical pastures.     

Post-dispersal seed predation of three grassland species in a plant diversity experiment

Aims Post-dispersal seed predation is an important ecosystem process because it can influence the seed's fate after the initial dispersal from the mother plant and subsequently transform communities. Even at small scales, post-dispersal seed predation can vary greatly depending on seed identity, granivorous taxa or microhabitat structure. However, little is known about the role of plant species richness and functional group richness in post-dispersal seed predation. The overall aim of this study was to test whether increasing plant species richness or plant functional group richness affects the rate and variability of post-dispersal seed predation. We additionally investigated the influence of vegetation structure and seed species identity on the rate and variability of post-dispersal seed predation and whether the influence of different granivorous taxa changed with increasing plant species richness.Methods We conducted seed removal experiments along a long-term experimental plant diversity gradient, comprising plots with monocultures to 60 species mixtures of common grassland species in Jena, Germany, in August 2011. We studied seeds of Onobrychis viciifolia, Pastinaca sativa and Trifolium pratense in exclusion experiments (seed cafeterias), an experimental setup that allowed access either for arthropods or slugs or for all granivorous taxa. Traditionally, seeds removed from seed cafeterias were classified as consumed but we used traceable fluorescent-coloured seeds to obtain more accurate predation rates by subtracting recovered seeds from overall removed seeds. The effect of multiple vegetation variables on mean and variability of seed predation rates was analysed using generalized mixed-effect models and linear regressions, respectively.Important findings Rates of recovered seeds were low but contributed to significant differences between seed predation rates and removal rates of seeds in some treatments. Seed predation rates were not directly correlated with increasing plant species richness or plant functional group richness but were influenced byseed species identity and granivorous taxa. Vegetation variables such as vegetation height and cover were significantly associated with seed predation rates. Depending on the seed species and/or the granivorous taxa, different vegetation variables correlated with seed predation rates. Our results indicate that effects of plant functional group richness and multiple vegetation variables on the magnitude of post-dispersal seed predation varied with seed identity and seed predator taxa. A direct effect of plant species and plant functional group richness could be shown on the variability of post-dispersal seed predation for some seed species and their respective predators. Thus, the changes in magnitude of post-dispersal seed predation with increasing plant species richness could potentially impact the fitness of some plant species and thereby influence plant community structure.     

Post-dispersal seed predation and the establishment of vertebrate dispersed plants in Mediterranean scrublands

The post-dispersal fate of seeds and fruit (diaspores) of three vertebrate-dispersed trees, Crataegus monogyna, Prunus mahaleb and Taxus baccata, was studied in the Andalusian highlands, south-eastern Spain. Exclosures were used to quantify separately the impact of vertebrates and invertebrates on seed removal in relation to diaspore density and microhabitat. The three plant species showed marked differences in the percentage of diaspores removed, ranging from only 5% for C. monogyna to 87% for T. baccata. Although chaffinches (Fringilla coelebs) fed on diaspores, rodents (Apodemus sylvaticus) were the main vertebrate removers of seed and fruit. Two species of ant (Cataglyphis velox and Aphaenogaster iberica) were the only invertebrates observed to remove diaspores. However, the impact of ants was strongly seasonal and they only removed P. mahaleb fruit to any significant extent. While removal of seed by rodents was equivalent to predation, ants were responsible for secondary dispersal. However, their role was limited to infrequent, small-scale redistribution of fruit in the vicinity of parent trees. Rodents and ants differed in their use of different microhabitats. Rodents foraged mostly beneath trees and low shrubs and avoided open areas while the reverse was true of ants. Thus, patterns of post-dispersal seed removal will be contigent on the relative abundance and distribution of ants and rodents. Studies which neglect to quantify separately the impacts of these two guilds of seed removers may fail to elucidate the mechanisms underlying patterns of post-dispersal seed removal. The coincidence of both increased seed deposition by the main avian dispersers (Turdus spp.) and increased seed predation with increasing vegetation height suggested that selection pressures other than post-dispersal seed predation shape the spatial pattern of seed dispersal. Rather than providing a means of escaping post-dispersal seed predators, dispersal appears to direct seeds to microhabitats most suitable for seedling survival. Nevertheless, the reliance of most vertebrate-dispersed trees on regeneration by seed and the absence of persistent soil seed banks imply that post-dispersal seed predators may exert a strong influence on the demography of the plants whose seeds they consume. Even where microsites are limited, the coincidence of the most suitable microhabitats for seedling establishment with those where seed predation is highest provide a means by which selective seed predators can influence community composition. Received: 19 August 1996 / Accepted: 25 January 1997