南方红豆杉的天然更新格局及食果鸟类对其种子的传播

2006年9-12月,在皖南仙寓山对南方红豆杉(Taxus chinensis var.mairei)天然更新种群的分布格局及食果鸟类对其种子的传播作用进行了研究.结果表明:天然更新种群集中分布于仙寓山海拔540 m以上面积约1.1 hm2的山坡上,与毛竹混生.共统计红豆杉1 331株,其中高度30 cm以下的幼苗1 167株,占总数的87.68%.该红豆杉种群为缓慢增长型种群,空间水平格局呈聚集分布.经过对结实母树的定点连续观察,记录到10种鸟类取食红豆杉种子,其中红嘴蓝鹊(Urocissa erythrorhyncha)、栗背短脚鹎(Hemixos castanonotus)和绿翅短脚鹎(Hypsipetes mcclellandii)为主要的取食者和传播者.食果鸟类吞入种子后,消化果肉(假种皮)后,种子随粪便排出而得以传播.野外调查中,在鸟类经常活动的地点发现带有红豆杉种子新鲜的鸟类粪便,每个含有1～2颗种子.未被鸟类取食的落果自然掉落于树冠下,假种皮被昆虫取食或腐烂,种子在母树下能正常萌发,但存活率几乎为零.因此,鸟类在该红豆杉天然种群形成中起到了重要作用,它们将种子传播至适宜萌发和生长的环境中.     

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     

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。     

食果鸟类对红楠种子的传播作用

肉质果植物通常依赖食果鸟类等取食果实后经消化道过程传播种子, 以完成种群的更新。红楠(Machilus thunbergii)是亚热带森林的代表性种类, 其果实具有依赖鸟类等动物取食后传播的特点。2012 年、2013 年在梅花山国家级自然保护区,研究了鸟类对红楠果实的取食及种子的传播作用。结果如下: 取食红楠种实的鸟类共计18 种, 其中整吞果实的鸟类12 种,啄取果肉的鸟类5 种, 啄食种子的鸟类1 种。整吞果实鸟类中4 种鹎科和2 种鸦科鸟类访问频次和取食量较大, 是主要的种子传播者。鸟类主要以呕吐方式传播种子。取食后初停栖点与母树距离及地面种子散布地点的分析结果表明, 鸟类可以远距离扩散种子。地面种实可被啮齿动物或蚁类再次捕食或搬运。研究表明, 红楠可以借助鸟类实现种子传播和种群更新。     

濒危乔木红豆杉不同更新阶段的生境特征比较

以福建梅花山自然保护区濒危乔木红豆杉(Taxus chinensis(Pilger)Rehd.)为对象,对红豆杉的种子排放、幼苗更新、幼树更新3个更新阶段的生境特征进行研究,并比较生态位的变化,以此评价红豆杉不同更新阶段的生境需求。结果显示:随着个体发育生长,红豆杉更新生境发生了明显转变,种子排放地与幼苗生境空间一致性较高,而它们与幼树生境存在明显的空间不一致性。从生态位角度来看,种子排放地和幼苗生境重叠指数≤0.5的仅有海拔和坡度2个因子,这说明幼苗的空间格局受到鸟类传播的强烈影响,鸟类传播所排放的种子能萌发并生长成幼苗。幼苗和幼树生境重叠指数≤0.5的因子有海拔、坡向、植被类型,说明在大尺度景观因子和庇护植被层面上,植物的更新需求出现了明显的不一致。研究结果表明红豆杉的更新生境存在明显的阶段变化,可能造成植物更新出现建成限制,使幼树阶段成为更新的瓶颈阶段。     

斑块生境中啮齿类和蚁类动物对南方红豆杉地面种子的搬运

南方红豆杉(Taxus chinensis)在我国南方地区多分布在村落附近的斑块生境中,成熟后的种子直接落在母树下或被鸟类等动物搬运至其他斑块中。林下地面种子会被地面活动的动物搬运,继而影响种子命运及种群更新。于2016年、2017年南方红豆杉果期,在浙江天目山一个红豆杉种群分布地(临安市桐坑村),采用野外种子摆放实验的方法研究了动物对地面种子的搬运情况。结果表明:啮齿类动物取食是南方红豆杉地面种子消失的主要原因,但种子消失率在斑块间及边缘生境中的差异较大。母树林和竹林是啮齿类动物的主要觅食生境,两种生境的动物取食率明显高于山核桃种植园。边缘生境成为啮齿类斑块间移动的通道,而非觅食场所。4种啮齿类动物中,淡腹松鼠(Callosciurus pygerythrus)在母树林斑块以外的生境中出现频率最高,而其他地面搬运者回避利用山核桃种植园,仅在母树林和竹林斑块中搬运南方红豆杉地面种子。可见,斑块生境中动物改变了南方红豆杉地面种子的空间沉积格局,进而影响植物种群的更新。     

福建梅花山自然保护区南方红豆杉母树特征对访问鸟类多样性的影响

在破碎化生境斑块中,植物特征影响着鸟类多样性格局,进而影响斑块的生态功能维系。南方红豆杉(Taxus chinensis)为国家Ⅰ级保护植物,在我国南方地区多集群分布在破碎化生境斑块中。为了解斑块生境中南方红豆杉母树特征对访问鸟类多样性的影响,于2018和2019年果期在福建梅花山保护区红豆杉生态园采用样线法和样点法结合,研究南方红豆杉结实量、郁闭度和高度等特征对访问鸟类多样性的影响。结果表明:在红豆杉生态园中,共有67种鸟类,其中45种鸟类为食果鸟类。不同年份间和不同斑块(竹林斑块和常绿阔叶林斑块)的鸟类群落组成和优势鸟类存在一定的差异。共有21种鸟类访问南方红豆杉母树,其多样性特点受母树特征的影响。一元线性回归模型表明,南方红豆杉母树结实量与访问母树的鸟类数量和种数呈正相关;母树郁闭度与访问母树的鸟类数量呈正相关。可见,斑块生境中南方红豆杉母树特征是影响鸟类多样性的主要因素之一,进而影响植物的种子更新。     

城市绿地中鸟类对海南蒲桃的取食和传播作用

海南蒲桃(Syzygium cumini)是亚热带地区城市绿地中常见的绿化植物, 其果实数量多、果期长, 可为鸟类提供大量食物资源。2020年6月—8月借助 Safari 10×26 变焦双筒望远镜, 采用焦点扫描法对访问海南蒲桃果实(种子)的鸟类行为进行观察, 详细记录鸟类的种类、取食基质、访问频次、取食时间、取食数量和取食方式等信息, 探讨鸟类在海南蒲桃种子传播及种群更新中的生态作用。结果表明: 成熟的海南蒲桃能吸引7种食果鸟类对其种子进行取食,其中白头鹎(Pycnonotus sinensis)、红耳鹎(Pycnonotus jocosus)、白喉红臀鹎(Pycnonotus aurigaster)和黄臀鹎(Pycnonotus xanthorrhous)4种鸟类以整吞的方式取食海南蒲桃的种子, 属于种子潜在传播者。不同种鸟类对海南蒲桃果实的取食频次间存在显著差异(t=4.310, df=6, P < 0.01), 平均访问只数、平均取食时间和平均取食量间存在极显著差异(P < 0.001)。食果鸟类的形态特征(体长、体重和嘴峰大小)与取食数量间均呈正相关。鸟类主要以呕吐的方式传播海南蒲桃的种子, 观察期间在鸟类取食后停歇的树木下方地面上常散落着较多表面洁净、无果肉包裹的裸露种子, 且有大量的海南蒲桃幼苗成功更新, 说明城市绿地中的植物是可以借助鸟类的取食和传播来实现其种群的更新。     

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.     

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。     

南京中山植物园秋冬季鸟类对植物种子的传播作用

1999年10月20日－2000年1月20日,在南京中山植物园内随机收集鸟粪样品160份,共分离出874粒结构完整的种子和果核、3块鞘翅目昆虫残体和1块鸟类羽毛残块。已鉴定出842粒种子和果核,分属于16科20属26种（变种）。在鸟粪样品中出现频率较高的种子依次分别属于冬青（Ilex purpurea)(22.22%)、圆柏（Sabina chinensis)(11.11%)、盐肤木（Rhus chinensis)(10.63%)、朴树（Celtis sinensis)(9．18％）、爬山虎（Parthenocissus tricuspidata)(7.73%)、龙柏（S.chinensis cv.kaizuca)(7.25%)等;种子数量相对较多的植物种类主要有冬青（23．52％）、盐肤木（16．15％）、圆柏（13．54％）、爬山虎（7．96％）、龙柏（7．96％）、小果蔷薇（Rosa cymosa)(5.34%)等。除经鸟粪传播外,鸟类还通过衔取果实以及在吞食果实后将种子呕出的方式传播种子。初步发芽试验表明,鸟粪样品中的爬山虎、盐肤木的种子,以及被鸟呕出的樟树（Cinnamomum camphora)、楝树（Melia azedarach)的种子均可发芽出苗。鸟类传播种子使南京中山植物园内樟树、冬青、海桐（Pittosporum tobira)和红豆杉（Taxus chinensis)等栽培树种成功地侵入到位于植物园北缘的虎山山坡黑松（Pinus thunbergii)、枫香（Liquidambar formosana)群落、以及植物园内山溪边的枫杨（Pterocarya stenoptera)、朴树群落等生境中。鸟类对种子的传播作用扩大了南京中山植物园内那些具有肉质果实、种子具有坚硬种皮或种子包被于坚硬果核中的植物种类的分布范围,促进了它们的自然更新。     

Fruit size, crop mass, and plant height explain differential fruit choice of primates and birds

Seed dispersal by animals is an important ecological process shaping plant regeneration. In general, seed dispersers are highly variable and often opportunistic in their fruit choice. Despite much research, the factors that can explain patterns of fruit consumption among different animal groups remain contentious. Here, we analysed the interactions between 81 animal species feeding on the fruits of 30 plant species in Kakamega Forest, Kenya, during 840 h of observations. Our aim was to determine whether plant characteristics, fruit morphology, fruit colours and/or fruit compounds such as water, sugar, phenols and tannins explained the relative importance of fruit consumption by the two most important consumer groups, primates and birds. We found significant differences in fruit choice between both groups. Primates fed on larger fruits and on higher trees that had larger fruit crops, whereas birds were observed feeding on smaller fruits and on smaller plants producing fewer fruits. Fruit colours did not differ between fruits consumed by primates and those consumed by birds. However, differences in the fruit choice among frugivorous birds were associated with differences in fruit colours. Smaller plants with smaller fruits produced red fruits which contrasted strongly with the background; these fruits were dispersed by a distinct set of bird species. The contents of water, sugar, phenols and tannins did not differ between fruits eaten by primates and those eaten by birds. Some phylogenetic patterns were apparent; primates fed preferentially on a phylogenetically restricted subsample of large plants with large fruits of the subclass Rosidae. We discuss why the observed primate dispersal syndrome is most likely explained by a process of ecological fitting.     

The role of frugivorous birds in fruit removal and seed germination of the invasive alien Cotoneaster franchetii in central Argentina

Many invasive plant species have fleshy fruits that are eaten by native frugivorous birds which disperse their seeds and may facilitate their germination, playing an important role in plant invasion success. The fleshy‐fruited shrub Cotoneaster franchetii (Rosaceae) is an important invasive alien in the mountainous regions of central Argentina. To determine the role of avian frugivorous in fruit removal of this species, we conducted a frugivore exclusion experiment including bagged and unbagged branches in 75 plants of C. franchetii. At the end of the dispersal period, we compared the percentage of missing fruits (removed by birds + naturally dropped) in unbagged branches with the percentage of naturally dropped fruits in bagged branches. To assess whether any mechanism acting on seeds during their passage through bird guts (de‐inhibition by pulp removal and/or seed scarification) affects seed germination of this species, we compared percentage and speed of germination among seeds obtained from faeces of the native frugivorous Turdus chiguanco, from manually de‐pulped fruits, and from intact fruits. The percentage of missing fruits per shrub in unbagged branches was significantly higher than the percentage of naturally dropped fruits in bagged branches, suggesting that frugivorous birds play an important role in fruit removal of C. franchetii in the study area. Seeds from bird faeces and from manually de‐pulped fruits germinated in higher percentage and faster than seeds from intact fruits. Germination percentage and speed of seeds from manually de‐pulped fruits were significantly higher than those of gut‐passed seeds. These results indicate that T. chiguanco increases and accelerates seed germination of C. franchetii through pulp removal, but not through seed scarification. Overall, our findings indicate that native frugivorous birds facilitate the dispersal and germination success of C. franchetii, likely playing an important role in its invasion throughout the mountainous region of central Argentina.     

Consequences of frugivore diversity for seed dispersal, seedling establishment and the spatial pattern of seedlings and trees

Many plants depend on frugivorous animals for the dispersal of their seeds. However, it is only poorly known whether regional differences in frugivore diversity have consequences for seed dispersal, seedling establishment, and the spatial distribution of seedlings and trees. This comparative study of seed dispersal investigated the consequences of regional differences in frugivore diversity for two tree species of the genus Commiphora. C. harveyi was studied in South Africa where avian frugivore diversity is high, C. guillaumini was studied in Madagascar where the avian frugivore community is depauperate. At both study sites, the percentages of handled and dispersed seeds in Commiphora trees were quantified by fruit traps, and visitation rates, seed handling rates and dispersal rates were quantified for each animal species for two consecutive years. Seedlings were mapped and the spatial distribution of trees quantified. At both study sites, fruits were mainly eaten by birds. The total percentage of dispersed seeds in South Africa was significantly higher than in Madagascar (70.8% vs. 7.9%) because there was a lack of effective dispersers that swallowed seeds in Madagascar. Seed dispersal benefit, i.e. the increase in the probability of becoming established as a seedling away from parent trees due to dispersal was much higher in Madagascar (80 times higher probability) compared to South Africa (6 times higher). Corresponding with the different dispersal percentages, seedlings in South Africa were found at relatively large distances from the nearest Commiphora tree (median distance=21.0 m), whereas in Madagascar seedlings were found mostly under and close to the nearest Commiphora tree (median distance=0.9 m). Finally, Commiphora trees in the Malagasy study site were clumped, but were more randomly distributed in the South African study site. These results suggest that regional differences in frugivore diversity and behaviour strongly affect seed dispersal of trees, seedling establishment and the spatial distribution of seedlings and trees.     

Fruit color and fruit size of bird-disseminated plants in Japan

Fruit color, fruit size and phenology of bird-disseminated plants were examined in different climatic zones of Japan and the relationships between the plants and frugivorous birds were disscussed.Black fruit was the most common in warm-temperate areas and red was the most common in cool-temperate and subarctic zones. Red was more dominant in the lower layer of the forests in all climatic zones. Bicolor fruits were frequent in trees and were not found in herbs. Both in warm- and cool-temperate zones conspicuous fruits which are red and bicolored display were more frequent in summer than in winter.The diameter of most fruits were 4–11 mm. Fruits in warm-temperate were somewhat bigger than those in cooltemperate zone. In forest strata the fruits of shrubs were smaller than those of trees and herbs in width. However I found no relationships between fruit size and fruit color.The frugivorous birds could have influenced not only the evolution of seasonal differences in the proportion of fruit color between warm-temperate and cool-temperate region, but also affect the fruit size.     

Phytomelatonin in the leaves and fruits of wild perennial plants

Phytomelatonin has been documented in numerous flowering plants, mostly in cultivated species consumed by humans. Although frugivorous animals feed on fruits, the phytomelatonin content of these organs has hardly ever been tested in wild plants. The aim of this study was to determine the levels of phytomelatonin in the leaves and fleshy fruits of 31 wild perennial species known to be eaten by herbivorous and frugivorous mammals and birds. Considerable levels of phytomelatonin were found in the leaves of all the tested species, and some contained melatonin in their fruits as well. The melatonin content was found to vary significantly in different life forms (trees, shrubs, and climbers), with trees possessing the highest levels. The analysis revealed a significant positive correlation between the phytomelatonin levels in the leaves and the fruits of various species. However, the concentration found in the fruits was generally lower than that found in the leaves of the same species. Despite the presence of phytomelatonin in the fleshy fruits of different families, there was no noticeable common attribute among them. Phytomelatonin was exhibited in both the seeds and the pulp, with no obvious preference for either one. Although it was determined that ingested melatonin enters the bloodstream of birds and mammals, its specific role is still not certain. The potential impact of edible phytomelatonin on the circadian rhythm of herbivores and frugivores is discussed on the basis of these findings.     

Interactions between birds and fruit in a temperate woodland

Summary The phenology of fruit trees and avian consumption of fruit were examined in Wytham wood, Oxford in 1979–1980. Ripe fruit was available to and fed upon by birds from late August until early May. Observations made on a daily transect indicated that most of the fruit was eaten by tits and thrushes, but the two families differed in the seasonal consumption and species of fruit taken. Tits took fruits of Elder, Bramble, White Bryony, Honeysuckle, Black Bryony, and Woody Nightshade at the beginning of the season only, while thrushes consumed fruits of Elder, Bramble, Hawthorn, Sloe, Rose, and Ivy mainly in the middle and end of the season. Members of these two families also visited different habitat types following the consumption of fruit, probably effecting different patterns of dispersal. There was no correlation between feeding preferences and the abundance or profitabilities (as defined by Pyke et al. 1977) of fruits with respect to total nitrogen, total proteins, total carbohydrates, total fats, or kilocalories. It is concluded that other factors such as palatability or content of other nutrients may be important in determining the feeding preferences of different species of frugivorous birds.     

Removal of vertebrate-dispersed fruits in vegetation on fertile and infertile soils

Summary The hypothesis that more plant species with vertebrate-dispersed fruits occur on fertile soils because there is a greater probability of fruit removal from the parent plant was tested at 16 sites around Sydney, Australia. Removal rates from artificial fruit spikes were two and a half times greater on fertile than infertile soil sites, although this was not quite statistically significant. High variability in removal rate between sites was evident irrespective of fertility. Most removal occurred during the day indicating that birds were important consumers, rather than nocturnal mammals. Bird abundance and diversity did not differ between soil types. More frugivorous species were found in plant communities growing on fertile soil. Two models could explain the patterns observed. Firstly, plants with vertebrate-dispersed fruits could be favoured on fertile soils because of a high abundance of frugivorous birds accomplishing seed dispersal. Alternatively, plants with vertebrate-dispersed fruits could be favoured on fertile soil sites for some other reason and frugivorous birds could be attracted to these areas of abundant food. The correlation between soil fertility and the percentage of vertebrate-dispersed fruits was stronger than the correlation between soil fertility and removal rates and suggests that the second model is more likely to be true. Frugivorous birds are unlikely to be responsible for the high percentage of species with vertebrate-dispersed fruits in fertile soil environments.     

Scale‐dependent effects of forest restoration on Neotropical fruit bats

Neotropical fruit bats (family Phyllostomidae) facilitate forest regeneration on degraded lands by dispersing shrub and tree seeds. Accordingly, if fruit bats can be attracted to restoration sites, seed dispersal could be enhanced. We surveyed bat communities at 10 sites in southern Costa Rica to evaluate whether restoration treatments attracted more fruit bats if trees were planted on degraded farmlands in plantations or island configurations versus natural regeneration. We also compared the relative influence of tree cover at local and landscape spatial scales on bat abundances. We captured 68% more fruit bat individuals in tree plantations as in controls, whereas tree island plots were intermediate. Bat activity also responded to landscape tree cover within a 200‐m radius of restoration plots, with greater abundance but lower species richness in deforested landscapes. Fruit bat captures in controls and tree island plots declined with increasing landscape tree cover, but captures in plantations were relatively constant. Individual species responded differentially to tree cover measured at different spatial scales. We attribute restoration effects primarily to habitat structure rather than food resources because no planted trees produced fruits regularly eaten by bats. The magnitude of tree planting effects on fruit bats was less than previous studies have found for frugivorous birds, suggesting that bats may play a particularly important role in dispersing seeds in heavily deforested and naturally regenerating areas. Nonetheless, our results show that larger tree plantations in more intact landscapes are more likely to attract diverse fruit bats, potentially enhancing seed dispersal.     

Functional heterogeneity in a plant–frugivore assemblage enhances seed dispersal resilience to habitat loss

The ability of ecosystems to maintain their functions after disturbance (ecological resilience) depends on heterogeneity in the functional capabilities among species within assemblages. Functional heterogeneity may affect resilience by determining multiplicity between species in the provision of functions (redundancy) and complementarity between species in their ability to respond to disturbances (response diversity), but also by promoting the maintenance of biological information that enables ecosystems to reorganize themselves (ecological memory). Here, we assess the role of the components of the functional heterogeneity of a plant–frugivore assemblage on the resilience of seed dispersal to habitat loss. For three years, we quantified the distributions of fruits, frugivorous thrushes (Turdus spp.) and dispersed seeds, as well as frugivore diet and movement, along a gradient of forest cover in N Spain. The abundances and the spatial distributions of fruits and birds varied between years. The different thrushes showed similar diets but differed in spatial behavior and response to habitat loss, suggesting the occurrence of both functional redundancy and response diversity. Forest cover and fruit availability affected the spatial distribution of the whole frugivore assemblage. Fruit tracking was stronger in years when fruits were scarcer but more widespread across the whole fragmented landscape, entailing larger proportions of seeds dispersed to areas of low forest cover and open microhabitats. Rather than depending on redundancy and/or response diversity, seed dispersal resilience mostly emerged from the ecological memory conferred by the inter‐annual variability in fruit production and the ability of thrushes to track fruit resources across the fragmented landscape. Ecological memory also derived from the interaction of plants and frugivores as source organisms (trees in undisturbed forest), mobile links (birds able to disperse seeds into the disturbed habitat), and biological legacies (remnant trees and small forest patches offering scattered fruit resources across the landscape).