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.     

The influence of pulp lipids on fruit preference by birds

Use patterns of lipid-rich, fleshy fruits were examined for frugivorous birds. Approximately 20% of fleshy-fruited species within plant communities bear fruit with >10% dry weight lipids. These fruits are fed upon by the most heavily frugivorous bird species in those communities studied and are the focus of highly specialized relationships among birds and fruits. In aviary tests some temperate frugivores show preference for higher lipid native fruits and prefer higher-lipid artificial fruits with as little as 3% wet weight difference in lipid content. The importance of lipids in fruit preference by birds, and the possible effects of differing digestive physiologies in birds are discussed.     

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.     

Frugivory and seed dispersal by vertebrates in the Oriental (Indomalayan) Region

Current knowledge of frugivory and seed dispersal by vertebrates in the Oriental Region is summarized. Some degree of frugivory has been reported for many fish and reptile species, almost half the genera of non-marine mammals and more than 40% of bird genera in the region. Highly frugivorous species, for which fruit dominates the diet for at least part of the year, occur in at least two families of reptiles, 12 families of mammals and 17 families of birds. Predation on seeds in fleshy fruits is much less widespread taxonomically: the major seed predators are colobine monkeys and rodents among the mammals, and parrots, some pigeons, and finches among the birds. Most seeds in the Oriental Region, except near its northern margins, are dispersed by vertebrate families which are endemic to the region or to the Old World. Small fruits and large, soft fruits with many small seeds are consumed by a wide range of potential seed dispersal agents, including species which thrive in small forest fragments and degraded landscapes. Larger, bigger-seeded fruits are consumed by progressively fewer dispersers, and the largest depend on a few species of mammals and birds which are highly vulnerable to hunting, fragmentation and habitat loss.     

Fleshy fruit characteristics in a temperate deciduous forest of Japan: how unique are they?

This study investigated the fleshy fruit characteristics of 28 woody species in a Japanese temperate forest where large sedentary seed-dispersing mammals are present. We tested whether the findings in previous studies in temperate forests of Europe and North America are universal or not. Results have suggested that fruits of all species were eaten both by birds and mammals except for four species with larger fruits, which were eaten only by mammals. A gradient was found from a syndrome characterized by small, oily, and large-seeded fruits to a syndrome characterized by large, succulent, non-oily, and small-seeded fruits. The sizes and colors of the fruits were not conspicuously different from previous findings in Europe and North America. On the other hand, nitrogen and lipids in the fleshy part did not show seasonally increasing trends, or even seasonally decreasing trends in terms of dry weight. This result, suggesting the absence of community-level adaptation of fruit traits to migratory bird dispersers, contrasted with findings in Europe and North America. Large sedentary arboreal or tree-climbing mammals may have a greater effect on the evolution of fruit-disperser relations than opportunistic migratory birds.     

Phytomelatonin: a potential phytotherapeutic intervention on COVID-19-exposed individuals

Phytomelatonin is a pleiotropic molecule that originated in higher plants with many diverse actions and is primarily an antioxidant. The recent identification and advancement of phytomelatonin unraveled the potential of this modulatory molecule being considered a new plant hormone, suggesting its relevance in treating respiratory infections, including COVID-19. Besides, this molecule is also involved in multiple hormonal, physiological, and biological processes at different levels of cell organization and has been marked for its ability to cross the blood-brain barrier and prominent antioxidant effects, reducing mitochondrial electron leakage, up-regulating antioxidant enzymes, acting as a free radical scavenger, and interfering with pro-inflammatory signaling pathways as seen in mood swings, body temperature, sleep, cancer, cardiac rhythms, and immunological regulation modulators. However, due to its diversity, availability, affordability, convenience, and high safety profile, phytomelatonin has also been suggested as a natural adjuvant. This review discussed the origin, content in various plant species, processes of extraction, and detection and therapeutic potentials of phytomelatonin in treating COVID-19-exposed individuals.     

Linking structure and functionality in mutualistic networks: do core frugivores disperse more seeds than peripheral species?

Seed dispersal networks are often organized in nested structures in a way that a few core species can disproportionally affect the remaining species in a network, in both the ecological and evolutionary sense. Yet, the relative importance of core and peripheral species has not been properly tested in seed dispersal network studies. We determined core species from 10 local seed dispersal networks composed by fleshy‐fruited plants and frugivorous birds. Each of those local quantitative networks was characterized with a core quality value, a core score for each species and a threshold value between core and peripheral species. From a total of 52 bird and 69 plant species that interacted in the study area, only 8 and 15, respectively, were identified as core. Each local network had a core that comprised a small number of birds and plants, always lower than 30% of the interacting species. There was no difference in the quantitative component of seed dispersal effectiveness (QC) provided by the frugivorous bird assemblage to plant functional groups clustered according to their growth form and fruit characteristics. Core birds dispersed seeds from each of these plant functional groups with a higher QC than peripheral species. Thus, we empirically demonstrate for the first time that seed dispersal networks at a regional scale have a small core set of fruit‐eating birds, upon which heavily rely most fleshy‐fruited plants for their seed removal. Hence, the activity of just a few core frugivores could deeply impact the demography of an entire assemblage of fleshy‐fruited plants.     

Ornithochory and plant succession in mediterranean abandoned orchards

Data on the occurrence of species with fleshy diaspores and on breeding birds were collected in three abandoned orchards, resp. 8, 11 and 24 years after abandonment. Most of the 41 phanerophytes and vines with fleshy diaspores are also found in the borders of the orchards, the more so if the number of years since abandonment increases. Most species have small seeds and red or black coloured diaspores, most of which ripen in autumn. The frequent species show a characteristic distribution pattern in relation to the distance of the fruit trees: high densities near the trunk and uniform decrease with distance. This is explained by the behaviour of frugivorous birds. No correlation was found between distribution patterns and soil conditions. Age structure of colonizing species shows a distribution conforming an inversed J curve in the more recently abandoned orchard. The main conclusions are:

1. Pioncor trees are attractive for frugivorous birds and may act as nuclei. This supports the facilitation model.
2. The distribution of diaspores by birds helps to homogenize the species distribution on the regional level but at the site level individual differences in nucleation cause a heterogeneity.

     

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.     

EVOLUTIONARY ASPECTS OF FRUIT-EATING BY BIRDS

In spite of a considerable literature on fruit-eating, the general evolutionary implications of fruit as a source of food for birds have been neglected. A preliminary attempt is made to explore the evolutionary and ecological consequences of fruit-eating, considered as a mutual interaction between parent plant and dispersal agent. The relationship considered is that obtaining between fleshy fruits and the “legitimate” fruit-eating birds which digest the fleshy part of the fruit and void the seed intact. Evolutionary aspects of seed-eating are also briefly discussed. The “strategies” adopted by fruits for dispersal by birds result in the production of abundant food supplies which are easy of access and exploitable by many species of birds. By contrast, the predation of birds on insects leads to a heterogeneous, sparse and cryptic food supply, to exploit which many different hunting techniques are necessary. Two important evolutionary developments in birds are attributed to these differences in food supply: there tend to be more species in families of insectivorous than of frugivorous birds, and lek behaviour in tropical forest has evolved in predominantly frugivorous birds. The seasonal succession of fruits in temperate latitudes is discussed, and contrasted with the situation in the tropics, using examples from Europe and Trinidad. In general, the succession of ripe fruits in Europe seems to be adapted to the seasonal shifts of the bird populations, and the more nutritious fruits tend to have a more southerly distribution and to ripen later than the more succulent fruits. In the tropics the distinction between nutritious and succulent fruits seems to be largely one of habitat. The constant succession of ripe fruits throughout the year in the tropics probably depends on competition for dispersal by frugivorous birds, which thus ensure the maintenance of their own food supply. This may be regarded as a symbiosis at the level of the ecosystem.     

Directed vertebrate-mediated seed dispersal of a fleshy-fruited amaryllid in a heterogenous habitat

Most plants with fleshy fruits have seeds that are ingested by animals, but a less well-understood mode of seed dispersal involves fleshy fruits containing seeds that are discarded by frugivorous animals because they are too large or toxic to be ingested. We studied the seed dispersal biology of Haemanthus deformis, an amaryllid lily species found in a mosaic of bush clumps in a grassland matrix in South Africa. We asked whether seed dispersal is directed in and among bush clumps and whether germination and survival are greater for seeds dispersed to bush clumps than for those dispersed into grassland. Using camera trapping, we found that fruits are consumed mainly by birds and rodents. The pulp was removed from the seeds which were then discarded without ingestion. While many seeds were dispersed close to the parent plant, most (c. 78.5%) were dispersed further than 1 m away from the parent plant. Longer distance dispersal resulted mainly from birds flying off with fruits in their bill or from rodents engaging in scatter-hoarding behavior. Seedling survival was most successful within bush clumps as compared to grasslands and shade was identified as a primary requirement for seedling survival. Seeds from which the fruit pulp had been removed germinated faster than those in intact fruits. Haemanthus deformis deploys a system of directed seed dispersal, whereby both birds and rodents contribute to the dispersal of seeds within patchy bush clumps that are favorable for seedling survival.     

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.     

Forest association and phenology of wild coffee in Kibale National Park, Uganda

A study on the forest association and phenology of wild coffee ( Coffea canephora Pierre) was conducted in Kibale forest, Uganda. Nested quadrats were used to enumerate tree species, including coffee and herbaceous plants associated with forest and coffee stands. A total of 150 coffee trees was marked along transects and monthly scans carried out to score for fruits, flowers, leaves and leaf insect damage. Pre- and post-dispersal predation levels and coffee yield estimates were made by examining fruits from trees, forest floor and seasonal fruit falls into demarcated plots. In the forest, wild coffee stands are associated with low-quality forest types in terms of timber species (about 10.5 canopy species/study site) and low stocking densities of trees ≥ 50 cm d.b.h. (average 38 trees ha⁻¹ for each site) and poor forest regeneration. In the forest, wild coffee reproductive phases overlap with ripening, coinciding with flower bud and flower production. The variable peak ripening season falls between November and April. The wild coffee yields are generally low (average of 3.5 intact fruits 16 m⁻² month⁻¹), with low insect fruit/seed damage (4–19%) but high levels of wastage due to monkeys, bats and birds.     

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.     

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     

Fruit size in wild olives: implications for avian seed dispersal

1. The response of frugivorous birds to an enlargement of fruit size, and the consequences for both birds and plants, are analysed for the interaction between avian seed dispersers and olives ( Olea europaea) .
2. The enlargement of fruit size promotes a shift in frugivorous birds' feeding behaviour, from swallowing fruits whole to pecking pieces of pulp. The relative frequency of olive consumption using each feeding behaviour was assessed by combining field data on frequency of appearance of olive pulp and seeds with data from laboratory trials.
3. Sardinian Warblers ( Sylvia melanocephala ) and European Robins ( Erithacus rubecula ) were mainly peckers both on cultivated and wild olives. Blackcaps ( Sylvia atricapilla ) consumed wild olives mainly by swallowing but consumed cultivated olives (larger than the wild ones) primarily by pecking. Song Thrushes ( Turdus philomelos ) were primarily swallowers of both types of fruits.
4. Laboratory trials with Song Thrushes, Blackcaps and European Robins showed that: (a) all were able to peck fruits; (b) fruit size determined a shift from swallowing to pecking, as pecking frequency increased with the enlargement of the fruit size; (c) all the species had an increased fruit handling failure rate when trying to swallow increasingly large fruits; and (d) from the birds' perspective, small shifts in fruit size may have important consequences on fruit profitability.
5. Pecking on olives turns the mutualistic fruit–frugivore interaction into a fruit-pulp predator interaction, thus arising a conflict between the plant and frugivorous birds.
6. This study shows that heavy dependence on fruit is not always simply related to seed dispersal. The same frugivorous bird species can act as a seed disperser or a pulp predator for the same plant species. The threshold between these roles is highly influenced by the ratio gape size/fruit size.     

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.     

Why do fleshy-fruit plants of the mediterranean scrub intercept fall-but not spring-passage of seed-dispersing migratory birds?

Summary Production of fleshy fruits by 8 tree and shrub species, and patterns of their utilization by 6 resident and 6 transient bird species, were assessed in an Israeli Mediterranean scrub. Whereas more than half the ripe crop of plants that fruit in one relatively short burst is not removed by birds, the greater part of the crop of species that fruit throughout a long period, is removed. Of two plant species that fruit simultaneously, the one with an inconspicuous fruit is utilized by a single, year-round resident frugivorous bird, whereas the species with bright fruits is utilized by several non-resident and omnivorous bird species. A further two simultaneously fruiting species differ in fat content and the color of their conspicuous ripe fruits; the low-fat fruit being taken by a resident species and the high-fat fruit by a transient congener, just prior to the desert-crossing portion of its Fall-passage.Though more migrants pass through in Spring than in Fall, none of the fleshy-fruit plants fruit in Spring, and most fruit during the Fall. Two non-exclusive explanations for this phenomenon are (a) Fall dispersal ensures immediate germination, with least exposure to desiccation and predation; (b) Spring transients approaching their breeding territories may either be reluctant to spend much time on feeding, or mostly require proteins, amply supplied by the Spring flush of insects, typical of the Mediterranean region. Fall transients approaching the desert require mostly fats and carbohydrates, supplied by fruits. Fruits attract birds easily in the Fall, when insects are scarce.     

ADAPTATIONS FOR A TWO-PHASE SEED DISPERSAL SYSTEM INVOLVING VERTEBRATES AND ANTS IN A HEMIEPIPHYTIC FIG (FICUS MICROCARPA: MORACEAE)

Figs (Ficus spp., Moraceae) are considered a classic example of plants with fleshy fruits adapted for seed dispersal by vertebrates, usually mammals or birds. Partially covering the endocarp of each individual drupelet of F. microcarpa is a fleshy, discrete lipid-containing exocarp that suggests adaptation for seed dispersal by ants. This structure is highly attractive to ants. F. microcarpa drupelets from which the fleshy exocarp was experimentally removed were much less likely to be transported by ants than those with this structure intact. The exocarps retained their attractiveness to ants and were not visibly altered following passage of the entire fruit through the gut of a frugivorous bird, the Indian Hill Mynah (Gracula religiosa). Germination percentage was not significantly affected by gut passage or exocarp removal. These results suggest that F. microcarpa has a two-stage seed dispersal system, in which primary dispersal by vertebrates is followed by secondary dispersal by ants. Dispersal aided by ants may be of significance in the biology of this exotic hemiepiphyte in southern Florida, where it is naturalized and appears to be spreading.     

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。