食果鸟白头鹎与接骨草的分布

作为种子的传播者 ,以果实为生的动物在所食植物的繁殖周期中起着关键作用。尽管动植物相互作用的研究自达尔文开始就引起了进化生物学家和生态学家的关注 ,但和传粉者与植物的相互作用的研究相比 ,种子传播者与植物相互作用的研究在 2 0世纪 70年代后才真正开始[1] 。植物学家     

浙江天目山自然保护区鸟类对美洲商陆种子的传播

为了解外来植物美洲商陆Phytolacca americana的入侵方式,2008年8月至2010年7月,在浙江省天目山国家级自然保护区内研究了鸟类对美洲商陆种子的传播作用.观察到白头鹎Pycnonotus sinensis、领雀嘴鹎Spizixos semitorques和红嘴蓝鹊Urocissa erythrorh...     

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。     

Fruits foraging patterns and seed dispersal effect of frugivorous birds on Hippophae rhamnoides sinensis

Behaviors of 18 species of birds eating fruits of Hippophae rhamnoides spp.sinensis were observed from September 2003 to March 2004.Their foraging patterns were found to be very different and Can be divided into five classes:(1)direct swallowing the fruits on crown of the shrubs and sometimes regurgitating seeds soon after;(2)carrying the fruits to their perching sites and swallowing;(3)pecking the fruits from the shrubs to the ground,eating pulp and seeds but leaving pericarp;(4)pecking through the pericarp,eating pulp and leaving pericarp and seeds;(5)pecking through the pericarp on the top of fruits,and only eating seeds.These foraging patterns have different effects on seed dispersal of H.rhamnoides spp.sinensis.The germination experiment of three groups of seeds(seeds from feces,dry fruits and extracted seeds from dry fruits)was carried out.Although ingestion processes of birds had some adverse effects on the seed germination of H.rhamnoides spp.sinensis,the seeds from feces still have a relatively higher germination ratio.H.rhamnoides spp.sinensis provides food to a variety of frugivorous birds.and the birds disperse its seeds.Thus,a mutually beneficial relationship between the bird and the seed is formed.     

鸟类取食中国沙棘果实的方式及其对种子的传播作用

2003年9月～2004年3月,定期观察取食中国沙棘果实的鸟类及其取食方式。共记录取食中国沙棘果实的鸟类18种,其取食果实的方式主要有：1)直接在树冠上吞食果实,有时候在吞食后将种子呕出;2)将果实从树上衔走后,在栖息处吞食或啄食;3)将果实啄落至地面,然后取食果肉和种子,留下果皮;4)啄破果皮,吸食果肉,留下果皮及种子;5)从顶端将果皮啄破后,仅取食里面的种子。不同的取食方式决定了它们对沙棘种子的传播作用不同。收集鸟粪便中的种子与完整的干果实及人工剥离果肉和内果皮的种子做萌发对比实验,结果表明,鸟类消化道过程对种子的萌发有一定影响,但萌发率仍较高。沙棘为多种鸟类提供食物,而鸟类则为沙棘传播种子,它们之间形成一种互利关系。     

藏雪鸡(Tetraogallus tibetanus)春夏季食性

藏雪鸡(retraogallus tibetanus)生活在青藏高原及其毗邻地区,是国家二级保护动物.2016年5-8月,在青海省尖扎县,采集藏雪鸡新鲜粪便及可能食用的植物,采用粪便显微分析法研究了藏雪鸡食性.结果表明:藏雪鸡能以各种植物器官为食,叶片是其食用的主要器官,茎次之,再次是花和果(种子),根极少;藏雪鸡所食...     

Do frugivorous birds concentrate their foraging activities on those alien plants with the most abundant and nutritious fruits in the South African Mediterranean-climate region?

Alien plants with abundant and nutritious fruits may compete more effectively for avian dispersal services than native shrubs. This premise was examined by comparing daily foraging activity (visitation frequency, foraging bird number and foraging period) by four frugivorous bird species of different size on fruits of two native and two alien shrub species co-occurring at four different sites. Also, the quantities of seeds consumed daily by each of the four birds species from fruits of the four shrub species were measured and compared with the numbers of fruits and seeds, and the mass of fruits present in the shrub canopies as well as with their fruit monosaccharide concentrations. The quantities of seed consumed daily by four different size bird species (Columba arquatrix, Colius striatus, Pycnonotus capensis, Zosterops pallidus subsp capensis) were positively correlated with the numbers of seeds per m² of canopy area and with fruit mass and fruit monosaccharide content per m² of canopy area, as well as with the monosaccharide concentration of individual fruits, except in the C. arquatrix (African olive pigeon). All four bird species displayed the highest daily visitation frequencies on fruits of the alien Solanum mauritianum which were more abundant and nutritious than fruits of the other alien Lantana camara and fruits the natives Olea europaea subsp africana and Chrysanthemoides monilifera. They also all consumed greater quantities of seed daily from fruits of the alien S. mauritianum than from fruits of the other shrub species. These results corroborate proposals that frugivorous birds concentrate their foraging activities on those alien plants with the most abundant and nutritious fruits.     

南京中山植物园鸟类对香樟 果实（种子）的取食

香樟(Cinnamomum camphora)是亚热带地区广泛分布的常绿阔叶树种,其果实数量多、果期长,可为鸟类提供大量食物资源。2018年11月至2019年2月,借助Safari 10×26变焦双筒望远镜,采用焦点扫描法对南京中山植物园内访问香樟果实(种子)的鸟类行为进行观察,详细记录鸟类的种类、取食基质、取食次数、取食时间、取食数量和取食方式等信息。研究期间有效记录达48d,累计记录到27种鸟类1021次取食香樟果实(种子)行为,存在2种取食基质(地面和树上)和3种取食方式(整吞果实、啄食果肉和取食种子)。T检验显示,不同鸟类对香樟果实(种子)的取食次数存在显著差异(t=3.096,df=26,P 0.01);不同月份间鸟类对香樟香樟果实(种子)的平均访问只数、平均取食次数、平均取食时间和取食数量均存在极显著差异(P0.001)。白头鹎(Pycnonotussinensis)、乌鸫(Turdusmendarinus)和灰喜鹊(Cyanopicacyanus)是访问次数及取食量最多的3种鸟类。单因素方差分析(One-way ANOVA)表明,上述这3种鸟类的取食数量存在极显著差异(F3, 598=25.219,P 0.001)。按照时间统计,11月份访问的鸟类种类和数量最多,1月份鸟类的取食次数、时间和数量最多。鸟类的形态特征(体重、体长和嘴峰长)与取食数量呈显著正相关,其中,体重和体长对取食数量的影响比嘴峰长的影响更显著,但嘴峰长影响鸟类对果实的取食方式。在鸟类常活动和停歇的树木下方地面往往散落着较多表面洁净、无果肉包裹的香樟种子,且园内人工林和自然更新林地可见许多香樟幼苗,表明鸟类对香樟具有潜在传播作用。     

Avian fruit preferences across a Puerto Rican forested landscape: pattern consistency and implications for seed removal

Avian fruit consumption may ensure plant reproductive success when frugivores show consistent preference patterns and effectively remove and disperse seeds. In this study we examined avian fruit preferences and their seed-removal services at five study sites in north-central Puerto Rico. At each site, we documented the diet of seven common fruit-eating avian species from February to September 1998. Using foraging observations and area-based estimates of fruit abundance, we examined preference patterns of birds. We found that 7 out of 68 fleshy-fruited plant species were responsible for most of the fruit diet of birds. Seventeen plant species were preferred and four of them were repeatedly preferred across several study sites and times by at least one avian species. Preferred plant species comprised a small percentage of fleshy fruits at each site (<15% in four out of five study sites), but showed extended phenology patterns. The quantity of seeds removed by frugivore species was not strictly related to preferences. Some frugivores showing no preference could effectively remove more seeds from plants at some locations than species exhibiting constancy in their patterns of preference. Only two frugivores, Euphonia musica and Vireo altiloquous, removed most of the seeds of plants for which they exhibited repeated preference across the landscape. Preference patterns, particularly those exhibiting consistency in space and time for plant species having prolonged fruiting periods, may have important mechanistic consequences for the persistence, succession, and regeneration of tropical plant communities.     

Bird Assemblage and Visitation Pattern at Fruiting Elmerrillia tsiampaca (Magnoliaceae) Trees in Papua New Guinea

Most tropical trees produce fleshy fruits that attract frugivores that disperse their seeds. Early demography and distribution for these tree species depend on the effects of frugivores and their behavior. Anthropogenic changes that affect frugivore communities could ultimately result in changes in tree distribution and population demography. We studied the frugivore assemblage at 38 fruiting Elmerrillia tsiampaca, a rain forest canopy tree species in Papua New Guinea. Elmerrillia tsiampaca is an important resource for frugivorous birds at our study site because it produces abundant lipid-rich fruits at a time of low fruit availability. We classified avian frugivores into functional disperser groups and quantified visitation rates and behavior at trees during 56 canopy and 35 ground observation periods. We tested predictions derived from other studies of plant–frugivore interactions with this little-studied frugivore assemblage in an undisturbed rain forest. Elmerrillia tsiampaca fruits were consumed by 26 bird species, but most seeds were removed by eight species. The most important visitors (Columbidae, Paradisaeidae and Rhyticeros plicatus) were of a larger size than predicted based on diaspore size. Columbidae efficiently exploited the structurally protected fruit, which was inconsistent with other studies in New Guinea where structurally protected fruits were predominantly consumed by Paradisaeidae. Birds vulnerable to predation foraged for short time periods, consistent with the hypothesis that predator avoidance enhances seed dispersal. We identified seven functional disperser groups, indicating there is little redundancy in disperser groups among the regular and frequent visitors to this tropical rain forest tree species.     

食果鸟类在红豆杉天然种群形成中的作用

2005年10月初至2006年2月,在南京中山植物园研究了红豆杉(Taxus chinensis)天然更新种群的分布格局,野外定点观察了鸟类对红豆杉种子的取食行为,记录了红豆杉落果的命运,分析了食果鸟类在红豆杉种群形成中的作用。结果表明:红豆杉天然更新种群集中分布于距母树群10m以外面积约0.5hm2的山坡上,全部红豆杉幼苗、幼树共213株,主要分布在中下坡;幼苗和幼树的空间水平格局符合聚集分布特征。"果实"成熟后,一部分被鸟类直接在树上取食,一部分掉落母树下地面。经过对结实母树的定点、连续观察,记录到4种鸟类取食红豆杉"果实",其中红嘴蓝鹊(Urocissa erythrorhyncha)和白头鹎(Pyc-nonotus sinensis)为主要的取食者和传播者。食果鸟类整吞"果实",消化果肉(假种皮)后,种子经消化道随粪便排出,以此过程传播种子。在取食完红豆杉"果实"后,食果鸟类多飞向坡上。坡上排水良好,有适量的荫蔽,生境适宜红豆杉种子萌发和幼苗的成长。因此,食果鸟类将种子传播至适宜萌发和生长的环境中,对该红豆杉种群的天然更新起主要作用。同时,鸟类传播对于红豆杉种群扩散、占据新的生境也有着特殊的意义。     

Reliability of macaques as seed dispersers

Seed dispersal is an ecological process crucial for forest regeneration and recruitment. To date, most studies on frugivore seed dispersal have used the seed dispersal effectiveness framework and have documented seed-handling mechanisms, dispersal distances and the effect of seed handling on germination. In contrast, there has been no exploration of “disperser reliability” which is essential to determine if a frugivore is an effective disperser only in particular regions/years/seasons or across a range of spatio-temporal scales. In this paper, we propose a practical framework to assess the spatial reliability of frugivores as seed dispersers. We suggest that a frugivore genus would be a reliable disperser of certain plant families/genera if: (a) fruits of these plant families/genera are represented in the diets of most of the species of that frugivore, (b) these are consumed by the frugivore genus across different kinds of habitats, and (c) these fruits feature among the yearly staples and preferred fruits in the diets of the frugivore genus. Using this framework, we reviewed frugivory by the genus Macaca across Asia to assess its spatial reliability as seed dispersers. We found that the macaques dispersed the seeds of 11 plant families and five plant genera including at least 82 species across habitats. Differences in fruit consumption/preference between different groups of macaques were driven by variation in plant community composition across habitats. We posit that it is essential to maintain viable populations of macaques across their range and keep human interventions at a minimum to ensure that they continue to reliably disperse the seeds of a broad range of plant species in the Anthropocene. We further suggest that this framework be used for assessing the spatial reliability of other taxonomic groups as seed dispersers.     

内蒙古天然草原不同放牧压力下亚洲小车蝗的食性及营养生态位（英文）

植物表皮蜡质中的饱和链烷作为内源指示剂广泛用于评价放牧家畜的食性和食量, 但用于天然草原蝗虫食性的评价研究较少。为了探讨天然草原蝗虫的食性及其生态位变化, 本研究以内蒙古天然草原为研究对象, 于2003年7-8月沿降水梯度选择3种典型植物群落（小针茅Stipa klemenzii、 羊草Leymus chinensis和大针茅Stipa grandis群落）, 在每个植物群落不同放牧压力下小区随机做20个植被样方, 样方内植物齐地面刈割, 测定其地上生物量和物种多样性, 取主要植物种测定其链烷模式, 同时采集放牧小区优势蝗虫种亚洲小车蝗Oedaleus asiaticus的粪便, 测定其链烷模式, 运用链烷技术评价蝗虫的食性及其营养生态位。结果表明： 不同植物群落中优势牧草种类及其比例不同, 其链烷模式存在种间差异, 链烷技术可以评价亚洲小车蝗的食性。亚洲小车蝗的食性在不同植物群落及不同放牧压力下存在显著的差异, 在羊草和大针茅群落中, 亚洲小车蝗是禾草采食者, 主要采食羊草和糙隐子草Cleistogenes squarrosa, 且与绵羊的营养生态位重叠指数较低, 分别为0.0619和0.0172; 在小针茅群落中亚洲小车蝗是杂类草采食者, 主要采食无芒隐子草Cleistogenes songorica、 猪毛菜Salsola collina和小针茅, 且与绵羊的营养生态位重叠指数较高, 达到0.1815。因此, 放牧不仅改变了群落的植物种类组成, 而且直接影响了亚洲小车蝗的食物组成, 二者对食物资源利用存在一定程度的竞争。     

Vertically stratified frugivore community composition and interaction frequency in a liana fruiting across forest strata

Vertical stratification is a key feature of tropical forests and structures plant–frugivore interactions. However, it is unclear whether vertical differences in plant-frugivore interactions are due to differences among strata in plant community composition or inherent preferences of frugivores for specific strata. To test this, we observed fruit removal of a diverse frugivore community on the liana Marcgravia longifolia in a Peruvian rain forest. Unlike most other plants, Marcgravia longifolia produces fruits across forest strata. This enabled us to study effects of vertical stratification on fruit removal without confounding effects of plant species and stratum. We found a high number of visits of a few frugivore species in the understorey and a low number of visits of many different frugivores in the canopy and midstorey. Whereas partial and opportunistic frugivores foraged across strata with differing frequencies, obligate frugivores were only found eating fruits in the higher strata. Avian frugivores foraging in the canopy were mainly large species with pointed wings, whereas under- and midstorey avian foragers were smaller with rounded wings. Our findings suggest a continuous shift in the frugivore community composition along the vertical gradient, from a few generalized frugivores in the understorey to a diverse set of specialized frugivores in the canopy. This shift in the frugivore community leads to correlated, reciprocal changes from specialized to generalized plant-frugivore interactions. Thus, we conclude that vertical niche differentiation between species in tropical forests persists even when food resources are available across strata. This highlights its role for promoting biodiversity and ecosystem functioning.     

Endozoochorous seed dispersal by sympatric mustelids, Martes melampus and Mustela itatsi, in western Tokyo, central Japan

We investigated seed dispersal by two sympatric mustelid species, the Japanese marten (Martes melampus) and Japanese weasel (Mustela itatsi), along an intercity forest path in western Tokyo, central Japan, from Jul 2007 to Jul 2008. We aimed to investigate the effect of food/habitat preference of these mustelids (martens are semi-arboreal frugivores while weasels are terrestrial carnivores) on their seed dispersal characteristics, which determine their efficacy as seed dispersers. In total, we analyzed 478 fecal samples collected from the two mustelids (N_marten = 381, N_weasel = 97). The proportions of feces containing seeds for martens and weasels were 81.4% and 55.7%, respectively. The number of plant species whose seeds were found within the feces were 28 and 17, respectively. Almost all seeds within feces of both mustelids were intact. The number of plant species whose seeds were found within a single fecal sample ranged from one to four, but no significant difference was detected between the two mustelids. However, marten feces contained a significantly greater number of seeds of most plant species as well as total number of seeds than did weasel feces. The numbers of plant species and seeds represented in marten feces varied seasonally, but those represented in weasel feces did not. Our findings suggest the possibility that both mustelids act in some ways as seed dispersers, although martens seem to disperse a greater diversity and total amount of seeds.     

Frugivore Behavioural Details Matter for Seed Dispersal: A Multi-Species Model for Cantabrian Thrushes and Trees

Animal movement and behaviour is fundamental for ecosystem functioning. The process of seed dispersal by frugivorous animals is a showcase for this paradigm since their behaviour shapes the spatial patterns of the earliest stage of plant regeneration. However, we still lack a general understanding of how intrinsic (frugivore and plant species traits) and extrinsic (landscape features) factors interact to determine how seeds of a given species are more likely to be deposited in some places more than in others. We develop a multi-species mechanistic model of seed dispersal based on frugivore behavioural responses to landscape heterogeneity. The model was fitted to data from three-years of spatially-explicit field observations on the behaviour of six frugivorous thrushes and the fruiting patterns of three fleshy-fruited trees in a secondary forest of the Cantabrian range (N Spain). With such model we explore how seed rain patterns arise from the interaction between animal behaviour and landscape heterogeneity. We show that different species of thrushes respond differently to landscape heterogeneity even though they belong to the same genus, and that provide complementary seed dispersal functions. Simulated seed rain patterns are only realistic when at least some landscape heterogeneity (forest cover and fruit abundance) is taken into account. The common and simple approach of re-sampling movement data to quantify seed dispersal produces biases in both the distance and the habitat at which seeds arrive. Movement behaviour not only affects dispersal distance and seed rain patterns but also can affect frugivore diet composition even if there is no built-in preference for fruiting species. In summary, the fate of seeds produced by a given plant species is strongly affected by both the composition of the frugivore assemblage and the landscape-scale context of the plant location, including the presence of fruits from other plants (from the same or different species).     

Do frugivores enhance germination success of plant species? An experimental approach

Frugivorous birds are among the most important consumers of fleshy fruits particularly in sub-tropical and tropical forest ecosystems. Whether or not such plant–frugivore interactions contribute to germination enhancement is still a subject of much debate. We tested the effect of gut treatment by four captive species of avian frugivores in comparison to manually depulped seeds and whole fruits on seedling emergence and germination probability of seeds from sixteen plant species in South Africa. Moreover, we determined whether fruit weight of each plant species affected germination patterns. Across plant species, a total of 2795 seeds were planted, of which 50% germinated. Both seedling emergence and germination probability neither differed among the bird species nor in comparison to manually depulped seeds or whole fruits. Further, seedling emergence and germination probability were both unaffected by fruit weight. However, the germination probability of all treatments increased similarly with increasing number of weeks after planting. Overall, these results suggest that seed depulping, neither by gut treatment nor manually improved germination of seeds, irrespective of their fruit weights. Thus, the major contribution of frugivores to forest regeneration may be more confined in transporting seeds away from the mother plant than in germination enhancement per se.     

Can network metrics predict vulnerability and species roles in bird‐dispersed plant communities? Not without behaviour

Network metrics are widely used to infer the roles of mutualistic animals in plant communities and to predict the effect of species' loss. However, their empirical validation is scarce. Here we parameterized a joint species model of frugivory and seed dispersal with bird movement and foraging data from tropical and temperate communities. With this model, we investigate the effect of frugivore loss on seed rain, and compare our predictions to those of standard coextinction models and network metrics. Topological coextinction models underestimated species loss after the removal of highly linked frugivores with unique foraging behaviours. Network metrics informed about changes in seed rain quantity after frugivore loss. However, changes in seed rain composition were only predicted by partner diversity. Nestedness, closeness, and d’ specialisation could not anticipate the effects of rearrangements in plant–frugivore communities following species loss. Accounting for behavioural differences among mutualists is critical to improve predictions from network models.     

Fruit–frugivore interactions in two southern hemisphere forests: allometry,phylogeny and body size

The size of fleshy fruits spans several orders of magnitude. However, the evolution of fruit size diversity is poorly understood. Fruit size diversity is hypothesised to result from several potential processes. The frugivore hypothesis postulates that different‐sized animal fruit consumers select for different‐sized fruits. The correlated selection hypothesis postulates that fruit size is allometrically related to other plant traits (e.g. leaf size, plant height); therefore differences in fruit size result from correlated evolution with other plant traits. We tested the frugivore and correlated selection hypotheses as potential explanations for fruit size diversity in two New Zealand study sites. We observed birds foraging for fruits over two fruiting seasons at each site and measured fruit size, leaf size and plant height in a total of 32 plant species. Relationships between average fruit size, leaf size, plant size and the average size of birds consuming each fruit species were then evaluated using phylogenetically independent contrasts. Similar results were obtained in both study sites. Fruit size was correlated with the size of avian fruit consumers, but was unrelated to leaf size or plant height. Therefore, results falsified the correlated selection hypothesis but failed to falsify the frugivore hypothesis. Although results suggest that frugivores may have influenced the evolution of fruit size in New Zealand, further study is needed to generate a mechanistic understanding of how frugivores may have selected for interspecific variation in fruit size.     

The role of frugivory in plant diversity maintenance – a simulation approach

Frugivores may play a key role in plant species coexistence by equalizing the species’ representation in the seed rain. Rare species may benefit from enhanced dispersal if frugivores prefer locally scarce fruits, or if rare plants are found in neighborhoods of high fruit density. Using a simulation model of frugivorous birds foraging on landscapes we tested if increased diversity in the seed rain could emerge from rare‐biased fruit selection, from the spatial configuration of plants, or both. In the absence of rare‐biased fruit selection, frugivores were not able to increase the diversity of the seed rain in any of our simulated landscapes. In contrast, when frugivores were attracted to locally scarce fruits, we found increased diversity in the seed rain whenever frugivore mobility across the landscape was high and plant species were well‐mixed in the fruiting neighborhoods. Irrespectively of the behavioral mechanism involved, landscape fragmentation lead to losses in diversity and species richness of simulated communities. In all simulations, density‐dependent mortality of dispersed seeds increased diversity in the community of seedlings. However, landscape homogenization at the scale of frugivores movements decreased the magnitude of this diversification effect. In summary, our study shows that frugivory has the potential to increase diversity in the seed rain when frugivores display rare‐biased fruit choices, provided that rare and common plants form heterogeneous neighborhoods. They also show that fragmentation is a major threat for diversity maintenance in the early‐regenerating community. Finally, they show that rarity confers advantages during regeneration only if it occurs at the scale of frugivores’ foraging decisions.