Host compatibility of the cloud forest mistletoe Psittacanthus schiedeanus (Loranthaceae) in Central Veracruz, Mexico

The consequences of the seed deposition of the parasitic mistletoe Psittacanthus schiedeanus were evaluated in a 32-mo study. We conducted a field seed inoculation experiment to determine variation in seed adhesion, seed germination, seedling establishment, and plant survival to reproduction among five host species and to evaluate whether these post-dispersal processes explain mistletoe prevalence and specificity at the regional scale. Seeds without an exocarp were inoculated onto branches of individuals of the five most common host species identified in nature in central Veracruz, México. Seed fate was monitored for 2 yr, at weekly intervals for the first 2 mo and at 2-mo intervals thereafter. The height and diameter of experimental host branches and canopy cover above them were measured to see if these factors affected mistletoe establishment. Significant differences in seed attachment and seed germination were found among host species. Fewer seeds remained attached on experimental branches of Quercus germana than those of Liquidambar styraciflua, Acacia pennatula, and Platanus mexicana. Although significant differences in seed germination were observed among species (significantly greater on A. pennatula), >70% of mistletoe seeds germinated within the first 5 wk on all host species. Towards the end of the inoculation experiment, more mistletoe seedlings survived, grew, and then flowered on Liquidambar styraciflua than on A. pennatula, P. mexicana, Q. germana, or Q. leiophylla. Host branch initial height and diameter did not affect seedling survival, but seedlings survived better on trees where the canopy was more open. Our results suggest that Liquidambar styraciflua is the most compatible host species with P. schiedeanus in central Veracruz. Not surprisingly, Liquidambar is by far the most common host tree for P. schiedeanus in this area as well. We suggest that the observed local specialization is a result of seed dispersal as consequence of bird foraging and territorial behavior, host abundance, and host species compatibility.     

Linking frugivore activity to early recruitment of a bird dispersed tree,Eugenia umbelliflora (Myrtaceae) in the Atlantic rainforest

Seed dispersal by animals is a complex process involving several distinct stages: fruit removal by frugivores, seed delivery in different microhabitats, seed germination, seedling establishment, and adult recruitment. Nevertheless, studies conducted until now have provided scarce information concerning the sequence of stages in a plant's life cycle in its entirety. The main objective of this study was to evaluate the immediate consequences of frugivore activity for Eugenia umbelliflora (Myrtaceae) early recruitment by measuring the relative importance of each fruit‐eating bird species on the establishment of new seedlings in scrub and low restinga vegetation in the Atlantic rainforest, Brazil. We conducted focal tree observations on E. umbelliflora trees recording birds' feeding behaviour and post‐feeding movements. We also recorded the fate of dispersed seeds in scrub and low restinga vegetation. We recorded 17 bird species interacting with fruits in 55 h of observation. Only 30% of the handled fruits were successfully removed. From 108 post flight movements of exit from the fruiting trees, 30.6% were to scrub and 69.4% to low restinga forest. Proportion of seed germination was higher in low restinga than in the scrub vegetation. Incorporating the probabilities of seeds' removal, deposition, and germination in both sites, we found that the relative importance of each frugivorous bird as seed dispersers varies largely among species. Turdus amaurochalinus and Turdus rufiventris were the best dispersers, together representing almost 12% probability of seed germination following removal. Our results show the importance of assessing the overall consequence of seed dispersal within the framework of disperser effectiveness, providing a more comprehensive and realistic evaluation of the relative importance of different seed dispersers on plant population dynamics.     

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.     

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     

Effects of seed dispersal on Juniperus communis recruitment on a Mediterranean mountain

Abstract. The recruitment of the relict shrub Juniperus communis on a mountain in SE Spain was studied during the period 1994–1998. The main objective was to determine both the quantitative and qualitative effects of bird dispersal on seedling establishment. Seed removal by birds, seed rain, post‐dispersal seed predation, germination, and seedling emergence and survival were analysed in different microhabitats. Birds removed 53 ‐ 89% of the seeds produced by plants. Seed rain was spatially irregular as most seeds accumulated near stones used by birds as perches and below mother plants while a few seeds were dropped in wet meadows and open ground areas. Post‐dispersal seed predation by rodents affected < 10% of dispersed seeds but varied significantly among microhabitats. Only 3.6 ‐ 5.5% of dispersed seeds appeared viable, as many seeds had aborted or showed wasp damage. Seeds germinated in the second and third springs after sowing, reaching a germination percentage of 36%. Seedling emergence was concentrated in wet meadows. Seedling mortality was high (75–80%), but significantly lower in wet meadows, the only microhabitat where seedlings could escape from summer drought, the main mortality cause. Seed abortion, germination and seedling mortality proved to be the main regeneration constraints of J. communis on Mediterranean mountains. Birds exerted a strong demographic effect, although their qualitative effect was limited by abiotic factors which caused the pattern of seed rain to differ from the final pattern of recruitment between microhabitats.     

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.     

Non-redundancy in seed dispersal and germination by native and introduced frugivorous birds: implications of invasive bird impact on native plant communities

Seed dispersal by vertebrate animals is important for the establishment of many fleshy-fruited plant species. Different frugivorous species can provide different seed dispersal services according to their specific dietary preferences as well as behaviour and body traits (e.g. body size and beak size of birds). Our aim was to study redundancies and complementarities in seed dispersal and germination between the two main native seed disperser birds and the introduced silver pheasant Lophura nycthemera in the temperate Patagonian forests. For this, we collected fresh droppings from the studied species and analyzed seed content. We conducted germination trials for four plant species common in bird droppings; two native species (Aristotelia chilensis and Rhaphithamnus spinosus) and two invasive non-native species (Rubus ulmifolius and Rosa rubiginosa). Both native frugivorous birds and the silver pheasant dispersed fruits of non- native fleshy-fruited plants, but their roles were non-redundant in terms of species dispersed and effect on seed germination. The silver pheasant dispersed a proportionally high number of non-native seeds, while native birds dispersed a high number of native seeds. In addition, the effect of gut treatment in seed germination differed between seed dispersers. Native birds promoted the germination for the two native plant species studied, while the silver pheasant promoted the germination of one non-native plant. This suggests that seed dispersal by the silver pheasant may contribute to the spread of some invasive fleshy-fruited plants in the ecosystems that otherwise would not be dispersed by any other bird. The understanding of redundancies and complementarities on seed dispersal and germination between native and introduced birds will allow improving the management of fleshy-fruited non-native plants.     

Differential passage time of mistletoe fruits through the gut of honeyeaters and flowerpeckers: effects on seedling establishment

McKey's (1975) hypothesis that avian dispersers with a specialized gut provide higher quality seed dispersal than unspecialized frugivores was tested using grey mistletoe (Amyema quandang) fruits, and captive mistletoebirds (Dicaeum hirundinaceum) and spinycheeked honeyeaters (Acanthagenys refogularis) in arid South Australia. Mistletoebirds have a specialized gut, unlike spiny-cheeked honeyeaters. The gut passage time of A. quandang fruits through mistletoebirds was 820±29 s (mean±SE, n=188), compared to 2434±36 s (n=436) for honeyeaters. The seeds defecated by both bird species were deployed on twigs of host trees. Despite the longer retention time of fruit in the gut of honeyeaters, the germination percentage of seeds defecated by mistletoebirds (85% of 485 seeds) and honeyeaters (81% of 485 seeds) did not differ significantly 1 week after deployment. However, after 5 months, a significantly greater proportion of seedlings had established from seeds passed by mistletoebirds (42.7%) than from seeds defecated by honeyeaters (31.1%). The data support the notion that the more gentle treatment of seeds in the gut of specialized dispersers translates into higher seedling establishment.     

啮齿动物的贮藏行为与植物种子的扩散

绝大多数啮齿动物一方面取食大量的植物种子和果实,另一方面通过其贮食行为将植物种子和果实搬运到远离母树的地点,即扩散,并将它们分散埋藏在落叶下或浅表的土层中,从而影响种子和果实的时空分布,最后导致幼苗在有利的条件下发生和建成,实现植物更新。啮齿动物与植物种子和果实之间已广泛形成了互惠或协同进化的关系。啮齿动物的贮食行为主要通过以下几个过程对植物种子和果实的扩散产生影响：选择、搬运和埋藏以及随后对种子和幼苗存活和死亡的影响等。本综述了啮齿动物对植物种子和果实贮藏的研究结果,以期为进一步开展啮齿动物的贮食行为在植物种子和果实的扩散中的作用的研究提供参考。     

The restricted seed rain of a mistletoe specialist

Specialist frugivores are the dominant consumers of mistletoe fruit in many regions and have been shown to intensify infections of host plants as a result of their rapid gut passage rates and dependence on existing infections. The role of specialist frugivores in long distance dispersal of mistletoe and establishment of new infections is unclear, and has not been explicitly evaluated previously. Here we critically examine the premise that specialists are the dominant dispersers by examining the role of an Australian mistletoe specialist (mistletoebird Dicaeum hirundinaceum Dicaeidae) in dispersing mistletoe (Amyema preissii Santalales: Loranthaceae) seeds beyond infected host stands. We use two primary lines of evidence – presence of birds using remote call recorders, and presence of dispersed seeds via surveys for defecated seeds on host branches. The observed and inferred movements of the mistletoebird were wholly restricted to habitat patches containing mistletoe, and this bird was not observed to transport seeds to nearby uninfected host stands within the study system. While mistletoe specialists may provide much of the within‐stand dispersal service for mistletoes, this serves only to aggregate and intensify existing infections. We suggest that long distance dispersal of mistletoe seeds beyond existing hosts and infection centres is not performed by these dietary specialists, these services more likely to be provided by generalist frugivores and other occasional mistletoe fruit consumers.     

Effectiveness of Mistletoe Seed Dispersal by Tyrant Flycatchers in a Mixed Andean Landscape

The dependence of mistletoes on few dispersers and the directed dispersal they provide is well known, yet no recent work has quantified either the effectiveness of these ‘legitimate’ dispersers, or the extent of redundancy among them. Here, I use the seed dispersal effectiveness (SDE) framework to analyze how birds (Mionectes striaticollis and Zimmerius bolivianus) contribute to mistletoe (Struthanthus acuminatus and Phthirusa retroflexa) infection in traditional mixed plantations within a humid montane forest in Bolivia. I calculated SDE for each bird–mistletoe pair and for the disperser assemblage, by estimating both the quantity and the quality of dispersal. The quantity of dispersal was measured as: (1) disperser abundance; (2) frequency of visits; and (3) number of seeds dispersed per visit, and the quality of dispersal was measured as: (1) germination percentage and speed of germination of seeds regurgitated by birds; and (2) the concordance of deposited seeds and seedling distribution patterns with adult mistletoe distribution at three scales (habitat, host, and microhabitat). Dispersers were not redundant: the more generalist species M. striaticollis dispersed more seeds, but provided lower quality seed dispersal, whereas the mistletoe specialist Z. bolivianus provided low‐quantity and high‐quality seed dispersal. Whereas S. acuminatus benefited more from the SDE of Z. bolivianus, P. retroflexa benefited from the complementary seed dispersal provided by both birds. These results demonstrate how sympatric mistletoes that share the same disperser assemblage may develop different relationships with specific vectors, and describe how the services provided by two different dispersers (one that provides high‐quality and one that provides high‐quantity dispersal) interact to shape spatial patterns of plants.     

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.     

Linking seed dispersal,germination and seedling recruitment in the invasive species <Emphasis Type="Italic">Berberis darwinii</Emphasis> (Darwin’s barberry)

Seedling recruitment is a multi-phased process involving seed production, dispersal, germination, seedling establishment and subsequent survival. Understanding the factors that determine success at each stage of this process is of particular interest to scientists and managers seeking to understand how invasive species spread and persist, and identify critical stages for management. To understand the factors and processes influencing recruitment of the invasive species Berberis darwinii Hook. (Darwin’s barberry), temporal and spatial patterns of seed dispersal, germination and seedling establishment were examined. Seed dispersal from a large source population was measured over two fruiting seasons, and subsequent patterns of seedling emergence and survival within each cohort were measured. Seed longevity was tested under both natural and artificial conditions. Seeds were widely dispersed by birds, up to 450 m from the source population. Dispersal was essential to seedling establishment, as few seedlings survived beneath the parent canopy. Seeds were relatively short-lived in the soil under both field and glasshouse conditions, with few surviving for more than 1 year. Patterns of newly emerged seedlings largely reflected patterns of seed rain, but seedling survival was significantly affected by distance from source population, seedling density and light environment. These results suggest that recruitment of B. darwinii is dependent on dispersal of seeds to favourable microsites. Management priorities should include the removal of fruiting plants, and seedling control in highlight areas.     

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

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

Are predatory birds effective secondary seed dispersers?

We have studied the unusual phenomenon of secondary seed dispersal of Lycium intricatum seeds on a small oceanic Atlantic island (Alegranza, Canarian Archipelago) in which a small frugivorous lizard ( Gallotia atlantica ) and two different predatory birds participate, a shrike ( Lanius excubitor ) and a kestrel ( Falco tinnunculus ). Endemic lizards that are common prey of both bird species consume Lycium fruits. Lizard remains were significantly matched with the presence of Lycium fruits in the regurgitation pellets of the two predatory birds. Seeds were found in 7.3% of the lizard droppings, 31.0% of kestrel pellets and 55.7% of shrike regurgitations. The mean number of seeds per dropping or pellet was 4.8 ±4 in lizard, 20.2 ±34.5 in shrike and 6.7 ±8.1 in kestrel. The percentage of viable seeds showed significant differences among all four treatments, decreasing in the following direction: seeds collected directly from plants (98.0%), shrikes (88.0%), lizards (72.3%), and kestrels (31.7%). Seeds from Lycium fresh fruits and shrike pellets showed significantly higher germination rates than those from lizard droppings and kestrel pellets. While lizards and shrikes are effective seed dispersers, kestrel gut treatment decreases seed viability. Seed viability is always higher than seed germination in each of the four treatments. In this island environment, Lycium seeds are under an important random influence during the seed dispersal process. Secondary seed dispersal seems to acquire a relevant dimension in small and remote insular environments or isolated continental systems where interactions among the different elements involved are intense, all of them are abundant native residents, and they have been coexisting for a long time. © 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002, 75 , 345–352.     

Generalist birds govern the seed dispersal of a parasitic plant with strong recruitment constraints

Mistletoes constitute instructive study cases with which to address the role of generalist consumers in the study of plant–animal interactions. Their ranges of safe sites for recruitment are among the most restricted of any plant; therefore, frugivores specializing in mistletoe have been considered almost indispensable for the seed dispersal of these parasitic plants. However, the absence of such specialists in numerous regions inhabited by many mistletoe species raises the question of whether unspecialized vectors may successfully disperse mistletoe seeds to narrowly defined safe sites. Using the European mistletoe Viscum album subsp. austriacum as a study case, we recorded a broad range of 11 bird species that disperse mistletoe seeds. For these species, we studied the mistletoe-visitation rate and feeding behavior to estimate the quantity component of dispersal effectiveness, and the post-foraging microhabitat use, seed handling, and recruitment probabilities of different microhabitats as a measure of the quality component of effectiveness. Both endozoochory and ectozoochory are valid dispersal mechanisms, as the seeds do not need to be ingested to germinate, increasing seed-dispersal versatility. Thrushes were the most effective dispersers, although they were rather inefficient, whereas small birds (both frugivores and non-frugivores) offered low-quantity but high-quality services for depositing seeds directly upon safe sites. As birds behave similarly on parasitized and non-parasitized hosts, and vectors have broad home ranges, reinfection within patches and the colonization of new patches are ensured by an ample assemblage of generalist birds. Thus, a parasitic plant requiring precision in seed dispersal can rely on unspecialized dispersers.     

Patterns in the evolution of amentiferous fruits

The diversity of fruit types within the Amentiferae appears to be the result of different seed dispersal and seedling establishment strategies that have evolved independently in the amentiferous families within the confines imposed by the wind-pollination syndrome. Dispersal of unadorned fruits and seeds by wind (e.g.,Betula, Rhoiptelea, Casuarina) has preceded the development of more efficient air-borne devices (e.g.,Ostrya, Carpinus, Engelhardia). Animal dispersal is the most advanced strategy, relying rarely on drupes, as in the shrubby Myricaceae, or on nuts, as inCorylus, Quercus, Carya, Juglans, Alfaroa, etc. The pattern of seedling establishment shows a structural-functional relationship to the seeddispersal strategy. Epigeal germination predominates in plants of open habitats. Those species with small fruits rely on wind dispersal, and their seeds have a relatively short germination time, whereas species with medium-sized fruits are often animal-dispersed with seeds that may have protracted germination times. In either case the reserve food supply for the young seedling is limited, and light is needed to spark photosynthesis. Hypogeal germination has evolved independently in several amentiferous families (e.g., Betulaceae, Fagaceae, Juglandaceae). This pattern is associated with closed habitats and plants with large seeds that have the capability of establishing vigorous seedlings in microhabitats of reduced light intensity where photosynthesis is impaired. The subterranean protection afforded the cotyledons and axillary meristems is a correlative feature that may have considerable importance in seedling survival where desiccation and predation are intense.     

Diplochory in western chokecherry: you can��t judge a fruit by its mesocarp

Western chokecherry (Prunus virginiana var. demissa, Rosaceae) is dispersed by frugivorous birds and carnivores, but it has large seeds that are potentially attractive to rodents that could act as seed predators and dispersers. Here, we quantify the benefits of primary dispersal by birds and secondary dispersal by scatter-hoarding rodents. In the fall, avian frugivores (mostly American robins, Turdus migratorius, and cedar waxwings, Bombycilla cedrorum) consumed 87% of the fruit crop and dispersed 67% of the fruit crop away from parent plants. Rodents removed 89% of seeds that simulated bird-dispersed seed rain from transects in riparian zones and 58% from transects in upland habitats. Rodents scatter-hoarded 91.6% of the seeds they removed, burying most in small caches (two to eight seeds) 8?C25?mm deep. About 39% of the seeds in spring caches produced seedlings. Inside rodent-proof exclosures, 52.1% of seeds buried to simulate rodent caches produced seedlings, 29.7% of which were still alive after 1?year. In contrast, only 3.8% of seeds placed on the soil surface, simulating dispersal by avian frugivores, produced seedlings. Seed dispersal by frugivorous birds likely contributes to colonization of unoccupied habitat through long-range dispersal and to escape from distance-dependent seed mortality near the parent plant. Despite seed losses, rodents offer short-range seed dispersal and bury seeds in more favorable sites for germination, improving seedling emergence and establishment. The combined mechanisms of seed dispersal significantly enhanced chokecherry seedling recruitment by providing more dispersal-related benefits than either frugivorous bird or scatter-hoarding rodents could provide alone.     

Spatial–temporal distribution of ornithochorous seeds from an Elaeagnus umbellata community dominating a riparian habitat

The dispersal efficiency and potential distribution of ornithochorous seeds of Elaeagnus umbellata in a riparian habitat were evaluated to clarify this species' establishment site in relation to the disturbance regime of the floodplain. Fruit removal by avian frugivores was monitored using fruit bags, and the spatial distribution of excreted seeds was quantified by seed traps set randomly on a gravel bar as an isolated seed source in the Yoshino River throughout an autumn fruiting season. Although more than 45% of the fruits remained on the twigs in the fruit bags, almost all fruits on the control twigs without fruit bags were exploited by the beginning of January. The fruit removal rate and seed dispersal distance were positively correlated with an increase in wintering bird species and their abundance. Intact bird‐dispersed seeds of E. umbellata were trapped within a 400‐m range and damaged seeds were limited to traps set within 50 m from the seed source. Frugivore behavior, such as feces excretion on rocks near water drinking sites and perching on surrounding woodland, greatly influenced the spatial and temporal dispersal pattern of the seed rain. In the present study, the avian frugivores showed upstream seed dispersal; thus, in years with stochastic autumnal floods, secondary dispersal via hydrochory downstream may be facilitated. The intensive seed dispersal in E. umbellata indicates that the present distribution of parent trees in the restricted elevation range of the gravel bars is the result of survival through disturbance, rather than seed dispersal limitation.     

Black bears Ursus americanus are effective seed dispersers,with a little help from their friends

Black bears Ursus americanus are generally considered effective seed dispersal agents for fleshy‐fruited plants because they can consume hundreds of fruits at once and have large home ranges. Although seedlings can emerge from faecal piles, establishment of such seedlings seems to be infrequent. Removal of seeds from faeces by rodents is often considered seed predation. We show that removal of seeds from bear faeces by seed‐caching rodents in the Sierra Nevada, USA, represents a second phase of seed dispersal that benefits some fleshy‐fruited plants. Using Trail Master infrared cameras to photograph animals and scandium‐46, a gamma‐emitting radionuclide, to track seeds, we determined that deer mice Peromyscus maniculatus removed seeds from bear faeces and cached them in soil. Caches typically contained 1–3 seeds buried 5–10 mm deep. These seeds escaped several sources of mortality by being moved to relatively safe locations, but deer mice also eventually eat many of the cached seeds. A field germination study confirmed that seed burial increased seedling emergence. Rodents removed seeds in bear faeces more quickly than those in bird faeces in one year, but seeds in bird faeces were removed faster in another year. Results varied across two years, probably because of availability of alternative food sources or changes in deer mice population sizes. The two‐phase seed dispersal syndrome described here may be important in understanding seed dispersal by carnivores and large ungulates that produce large faecal deposits containing many relatively large seeds.