Relationships between bird-dispersed plants and avian fruit consumers with different feeding strategies in Japan

We examined the interactions between bird-dispersed plants and fruit-consuming birds with various feeding strategies, by reviewing the plant species consumed by 14 bird species in Japan with four feeding types: gulpers (five species), grinders (four species), crushers (four species), and peckers (one species). Our literature review provided information on the plant species consumed by the birds in Japan and the morphological traits of the fruits: fruit volume, seed mass and number, pulp type (fleshy, dry, or arillate), and plant height (tall, medium, or small). Using these data, we examined the diversity of plant species consumed by each bird and the fruit morphological traits that affected fruit selection. The five gulpers consumed fruits from the largest number of plants, followed by the four grinders, the four crushers, and the one pecker. The gulpers and grinders consumed a wider variety of fruits than were consumed by the crushers and the pecker. Logistic regression analysis revealed that some crushers and the pecker preferred plants with dry or arillate pulp around the seeds. Our results suggest that a frugivorous bird’s feeding strategies, and particularly its fruit-handling behaviors and the fruit parts it ingests, influence the diversity of plants it consumes. The crushers and the pecker, which feed exclusively on seeds, require more effort and time to consume this type of food, and this might cause a strong preference for specific fruit traits and thus, consumption of a lower diversity of plant species.     

Crop size is more important than neighborhood fruit availability for fruit removal of <Emphasis Type="Italic">Eugenia uniflora</Emphasis> (Myrtaceae) by bird seed dispersers

For a plant with bird-dispersed seeds, the effectiveness of seed dispersal can change with fruit availability at scales ranging from individual plants to neighborhoods, and the scale at which frugivory patterns emerge may be specific for frugivorous species differing in their life-history and behavior. The authors explore the influence of multispecies fruit availability at two local spatial scales on fruit consumption of Eugenia uniflora trees for two functional groups of birds. The authors related visitation and fruit removal by fruit gulpers and pulp mashers to crop size and conspecific and heterospecific fruit abundance to assess the potential roles that facilitative or competitive interactions play on seed dispersal. The same fruiting scenario influenced fruit gulpers (legitimate seed dispersers) and pulp mashers (inefficient dispersers) in different ways. Visits and fruit removal by legitimate seed dispersers were positively related to crop size and slightly related to conspecific, but not to heterospecific fruit neighborhoods. Visits and fruit consumption by pulp mashers was not related to crop size and decreased with heterospecific fruit availability in neighborhoods; however, this might not result in competition for dispersers. The weak evidence for facilitative or competitive processes suggest that interaction of E. uniflora with seed dispersers may depend primarily on crop size or other plant’s attributes susceptible to selection. The results give limited support to the hypothesis that spatial patterns of fruit availability influence fruit consumption by birds, and highlight the importance of considering separately legitimate and inefficient dispersers to explain the mechanisms that lie behind spatial patterns of seed dispersal.     

Functional variation among frugivorous birds: implications for rainforest seed dispersal in a fragmented subtropical landscape

Seed dispersal plays a critical role in rainforest regeneration patterns, hence loss of avian seed dispersers in fragmented landscapes may disrupt forest regeneration dynamics. To predict whether or not a plant will be dispersed in fragmented forests, it is necessary to have information about frugivorous bird distribution and dietary composition. However, specific dietary information for frugivorous birds is often limited. In such cases, information on the seed-crushing behaviour, gape width and relative dietary dominance by fruit may be used to describe functional groups of bird species with respect to their potential to disperse similar seeds. We used this information to assess differences in the seed dispersal potential of frugivorous bird assemblages in a fragmented rainforest landscape of southeast Queensland, Australia. The relative abundance of frugivorous birds was surveyed in extensive, remnant and regrowth rainforest sites (16 replicates of each). Large-gaped birds with mixed diets and medium-gaped birds with fruit-dominated diets were usually less abundant in remnants and regrowth than in continuous forest. Small-gaped birds with mixed diets and birds with fruit as a minor dietary component were most abundant in regrowth. We recorded a similar number of seed-crushing birds and large-gaped birds with fruit-dominated diets across site types. Bird species that may have the greatest potential to disperse a large volume and wide variety of plants, including large-seeded plants, tended to be less abundant outside of extensive forests, although one species, the figbird Sphecotheres viridis, was much more abundant in these areas. The results suggest that the dispersal of certain plant taxa would be limited in this fragmented landscape, although the potential for the dispersal of large-seeded plants may remain, despite the loss of several large-gaped disperser species.     

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.     

Exclusive frugivory and seed dispersal of Rhamnus alaternus in the bird breeding season

We studied avian frugivory and seed dispersal in a dioecious shrub, Rhamnus alaternus, focusing on the quantitative and qualitative components of effectiveness. The study took place at three locations in the northeast of the Iberian Peninsula, and examined bird behaviour, intensity of feeding, and the consequences for seedling emergence. The coincidence between the bird breeding season and fruit ripening of R. alaternus in the absence of other ripe fruit, generates a monospecific interaction. The extant frugivorous species were mainly legitimate seed dispersers and their abundance was low. Sylvia melanocephala and S. undata were the most important at one site whereas S. atricapilla, Erithacus rubecula and Turdus merula predominated at the other two sites. Fruit handling took place directly on the branches. Bird species used microhabitats differently as first post-feeding perch, which usually was a short distance away. The low density of frugivorous birds in all localities, among others factors, resulted in satiation of the disperser community and many mature fruits unconsumed. Both adults and juveniles feed upon the plants and their foraging patterns are similar. Adults of S. melanocephala were observed to feed fruit to nestlings and consequently a second phase of dispersal potentially arises from the transport of fecal sacs. Pulp removal and passage through the digestive tract increased the probability of seedling emergence. This plant-dispersal interaction has important consequences, both positive and negative for the plant. Positively, the fruiting of R. alaternus at a time when other ripe fruits are not available avoids interspecific competition for seed dispersers. In addition, a low density of seed rain may reduce intraspecific competition. Negatively, the low density and small size of the breeding frugivorous bird community limit fruit handling and removal away from the parent plants, while the territorial behaviour of birds at that time of the year reduces the potential distances of seed dispersal.     

Frugivory,Post‐feeding Flights of Frugivorous Birds and the Movement of Seeds in a Brazilian Fragmented Landscape

Habitat fragmentation can break down the movement processes of frugivorous animals, thus influencing the relationship between plants and their seed dispersers by altering the number and identity of seed dispersers, and their relative contribution to seed dispersal. We studied the assemblages of frugivorous birds, their composition, species richness, and visitation rates to fruiting plants growing in the different landscape elements (forest fragments, live fences, and trees isolated in pastures) embedded in a Brazilian fragmented, agricultural landscape. By following the post‐feeding movements of frugivorous birds, we inferred the direction of seed movement from and to each of these landscape elements. Fruiting trees growing at different landscape elements were visited by frugivorous birds at similar rates. Isolated trees attracted a greater and distinct bird assemblage than trees in forest fragments or live fences. Judging by the post‐feeding flights of birds, the seeds of isolated trees were the most likely to reach all the landscape elements considered, but the contribution of isolated trees to the seeds falling in forested habitats or pastures depended on their degree of isolation. A few bird species were able to move widely, visiting fruiting plants in all landscape elements, and promoting long‐distance dispersal for plants. These few birds are of special interest because they are mobile links that connect habitats in fragmented landscapes with their seed dispersal services. Abstract in Spanish is available at http://www.blackwell‐synergy.com/loi/btp .     

Effects of site and fruit size on the composition of avian frugivore assemblages in a fragmented Afrotropical forest

Factors influencing the interaction between fruiting trees and their frugivorous seed dispersers in fragmented Afrotropical landscapes are poorly known. With the use of Mantel statistics we analysed assemblages of frugivorous birds on 58 individual trees belonging to 11 species growing in seven Kenyan cloud forest fragments. Overall, frugivores showed little specialization on particular trees. Fruit size explained a substantial amount of the variation in frugivore assemblages among different tree species at the same site. In addition, frugivore assemblages on conspecific trees were significantly more similar when the trees occurred at the same site. This location effect was attributable to the different sites and forest fragments (of different sizes and disturbance levels) varying in the densities and composition of their avian frugivores, vegetation composition and tree fruiting phenologies. It was consolidated further by the low mobility of most of these avian frugivores, particularly their reluctance to cross between forest fragments. Habitat disturbance and fragmentation may therefore have affected fruit selection, with implications for both seed dispersal and regeneration.     

Can Functional Traits Predict Ecological Interactions? A Case Study Using Rain forest Frugivores and Plants in Australia

In rain forest, the large numbers of species of fleshy-fruited plants and frugivorous animals result in a large number of potential fruit–frugivore interactions, which are challenging to survey in the field. Yet, knowledge of these relationships is needed to predict consequences of changes in the frugivore assemblage for seed dispersal. In the absence of comprehensive dietary information, it may be possible to delineate between frugivores that disperse different plants using ‘functional traits,’ or morphological and behavioral attributes of frugivores that interact with differences in salient characteristics of plant species. Here we use data on the consumption of 244 Australian rain forest plant species by 38 bird species to test for associations between patterns of frugivory and birds': (1) degree of frugivory, (2) gape width, and (3) seed treatment (seed crushing or seed dispersing). Degree of frugivory and gape width explain 74 percent of the variation in the sizes of fruits consumed by frugivorous birds. Among birds that consume a substantial dietary proportion of fruit, birds with wider gapes consume larger fruits. In contrast, this relationship was not shown by birds for which fruit is only a minor dietary component. Degree of frugivory and gape width, together with seed treatment, also strongly predict the overall taxonomic composition and diversity of plants consumed by bird species. Functional classifications of frugivore species may prove useful in developing a predictive understanding of fruit–frugivore interactions in other rain forest regions where detailed dietary information is not available for most frugivores.     

Bird attributes, plant characteristics, and seed dispersal of Pera glabrata (Schott, 1858), (Euphorbiaceae) in a disturbed cerrado area

Several plant characteristics, such as fruit production, nutrient reward, secondary compounds, and fruit color display, affect fruit choice by birds. On the other hand, several bird attributes affect their efficiency as dispersers. Here we investigate the ornithochoric seed dispersal of Pera glabrata Schott (Euphorbiaceae) in a cerrado fragment in southeastern Brazil. A set of bird attributes, such as frequency of visits, number of diaspores eaten, time spent foraging, methods of taking and handling the diaspores and agonistic interactions were analyzed in order to infer about the potential of each species to act as a seed disperser. Birds were the unique seed dispersers of these oil-rich diaspores. We observed 414 bird visits during 60 hours of focal observations in five trees from December 1999 to January 2000. Twenty bird species from seven families ate the diaspores of P. glabrata, but only 14 species were considered potential seed dispersers because they swallowed the diaspores, increasing the probabilities for the seeds to be defecated and/or regurgitated away from the parent trees. The main potential seed dispersers were: Turdus leucomelas (Muscicapidae), Dacnis cayana (Emberizidae), Colaptes melanochloros (Picidae) and Elaenia spp. (Tyrannidae). We did not find any significant seasonal change in the number of visits on the fruiting trees throughout the day. We also did not find any relation between the number of visits per tree and fruit production. The most effective seed dispersers of P. glabrata were generalist birds, which have a high visiting rate, high fruit consumption rate, and spend short periods on the plants. The large number of species recorded as potential seed dispersers of P. glabrata, being most of them very abundant even in Brazilian disturbed areas, may guarantee seed dispersal of this plant in small fragments and regenerating areas.     

Opposed latitudinal patterns of network‐derived and dietary specialization in avian plant–frugivore interaction systems

Latitudinal patterns of biodiversity have been studied for centuries, but it is only during the last decades that species interaction networks have been used to examine the proposed latitudinal gradient of biotic specialization. These studies have given idiosyncratic results, which may either be because of genuine biological differences between systems, different concepts and scales used to quantify biotic specialization or because the methodological approaches used to compare interaction networks were inappropriate. Here we carefully examine the latitudinal specialization gradient using a global dataset of avian plant–frugivore assemblages and interaction networks. In particular, we test whether network‐derived specialization patterns differ from patterns based on assemblage‐level information on avian dietary preferences on specific food types. We found that network‐derived measures of specialization (complementary specialization H₂′ and < d’>, modularity Q) increased with latitude, i.e. frugivorous birds divide the niche of fruiting plants most finely at high latitudes where they also formed more modular interaction networks than at tropical latitudes. However, the strength and significance of the relationship between specialization metrics and latitude was influenced by the methodological approach. On the other hand, assemblage‐level information on avian specialization on fruit diet (i.e. the proportion of obligate frugivorous bird species feeding primarily on fruit) revealed an opposed latitudinal pattern as more bird species were specialized on fruit diet in tropical than in temperate assemblages. This difference in the latitudinal specialization gradient reflects that obligate frugivores require a high diversity of fruit plants, as observed in tropical systems, and fulfil more generalized roles in plant–frugivore networks than bird species feeding on different food types. Future research should focus on revealing the underlying ecological, historical and evolutionary mechanisms shaping these patterns. Our results highlight the necessity of comparing different scales of biotic specialization for a better understanding of geographical patterns of specialization in resource–consumer interactions.     

Seed dispersal of fleshy-fruited invasive plants by birds: contributing factors and management options

The ecology of seed dispersal by vertebrates has been investigated extensively over recent decades, yet only limited research has been conducted on how suites of invasive plants and frugivorous birds interact. In this review, we examine how plant fruit traits (morphology, colour and display, nutritional quality, accessibility and phenology), avian traits (fruit handling techniques, gut passage time and effect, bird movements and social behaviour and dietary composition) and landscape structure (fruit neighbourhood, habitat loss and fragmentation and perch tree effects) affect frugivory and seed dispersal in invasive plants. This functional approach could be used to develop generic models of seed dispersal distributions for suites of invasive plant species and improve management efficiencies. Four broad research approaches are described that could direct management of bird‐dispersed invasive plants at the landscape scale, by manipulating dispersal. First, research is needed to quantify the effect of biological control agents on dispersal, particularly how changes in fruit production and/or quality affect fruit choice by frugivores, dispersal distributions of seed and post‐dispersal processes. Second, we explore how seed dispersal could be directed, such as by manipulating perch structures and/or vegetation density to attract frugivorous birds after they have been foraging on invasive plant fruits. Third, the major sources of seed spread could be identified and removed (i.e. targeting core or satellite infestations, particular habitats and creating barrier zones). Fourth, alternative food resources could be provided for frugivores, to replace fruits of invasive plants, and their use quantified.     

Fragmentation, habitat composition and the dispersal/predation balance in interactions between the Mediterranean myrtle and avian frugivores

Human‐induced fragmentation and disturbance of natural habitats can shift abundance and composition of frugivore assemblages, which may alter patterns of frugivory and seed dispersal. However, despite their relevance to the functioning of ecosystems, plant‐frugivore interactions in fragmented areas have been to date poorly studied. I investigated spatial variation of avian frugivore assemblages and fruit removal by dispersers and predators from Mediterranean myrtle shrubs (Myrtus communis) in relation to the degree of fragmentation and habitat features of nine woodland patches (72 plants). The study was conducted within the chronically fragmented landscape of the Guadalquivir Valley (SW Spain), characterized by ~1% of woodland cover. Results showed that the abundance and composition of the disperser guild was not affected by fragmentation, habitat features or geographical location. However, individual species and groups of resident/migrant birds responded differently: whereas resident dispersers were more abundant in large patches, wintering dispersers were more abundant in fruit‐rich patches. Predator abundances were similar between patches, although the guild composition shifted with fragmentation. The proportion of myrtle fruits consumed by dispersers and predators varied greatly between patches, but did not depend on bird abundances. The geographical location of patches determined the presence or absence of interactions between myrtles and seed predators (six predated and three non‐predated patches), a fact that greatly influenced fruit dispersal success. Moreover, predation rates were lower (and dispersal rates higher) in large patches with fruit‐poor heterospecific environments (i.e. dominated by myrtle). Predator satiation and a higher preference for heterospecific fruits by dispersers may explain these patterns. These results show that 1) the frugivore assemblage in warm Mediterranean lowlands is mostly composed of fragmentation‐tolerant species that respond differently to landscape changes; and 2) that the feeding behaviour of both dispersers and predators influenced by local fruit availability may be of great importance for interpreting patterns of frugivory throughout the study area.     

Resource tracking by avian frugivores in mountain habitats of northern Spain

Systems comprising avian frugivores and fleshy‐fruited plants are commonly used as models to study how animals respond to temporal and spatial variations in food abundance and distribution. Results have been far from conclusive, partly because of methodological constraints. In this study we investigated the fulfilment of a necessary condition for food tracking: correlation over time and space between the abundance of food and the abundance of food trackers. We used a paired‐sample design in which fruit sites were compared with fruitless sites on two different spatial scales (landscape and habitat patch) in eight different river basins in the Cantabrian Mountains (NW Spain). Fruit and bird abundances were recorded monthly (October to February) at all sites during three consecutive fruiting seasons. Fruiting seasons (“years”) could be ranked by fruit abundance as low (2003), intermediate (2002) and high (2004). Since fruit sites were located at considerably higher elevations than fruitless sites, the possible effects of altitude, together with those of other uninvestigated factors, were taken into account by using non‐frugivorous birds. Our results apparently suggest a temporal and spatial association between frugivores (avian seed dispersers) and fruit availability at the landscape scale but not at the smaller (habitat patch) scale. This, together with the absence of a similar pattern in non‐frugivores, suggests that fruit availability is an important ecological factor affecting the autumn and winter abundance patterns of frugivorous birds and that this factor operates at the landscape scale but not at the habitat patch scale. Moreover, the negative elevational gradient in bird abundance that is typical of temperate mountain ecosystems was clearly reversed for frugivores in the intermediate and high fruiting years (suggesting that the negative effect of higher altitude may have been counterbalanced by the greater availability of food). As in other European upland areas, traditional farming practices in the Cantabrian Mountains are declining and, as a result, the proportion of fruitless habitats is increasing. Our study suggests that in these upland ecosystems frugivorous bird abundances in autumn and winter are associated to fruit supply, thus this habitat change tendency will probably have, in the long‐term, a negative effect on bird populations.     

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.     

Avian translocations restore dual interaction networks in an island ecosystem

Many island bird species have been driven to extinction by introduced predators. Although poorly understood, these extinctions could have a 2-fold impact on bird–plant mutualisms, because island bird species can serve as both pollinators and seed dispersers. We investigated how avian translocations into a mammal-free reserve in New Zealand affected the structure of bird–flower and bird–fruit interactions. We observed bird–fruit and bird–flower interactions over a 9-year period to establish (1) the extent to which native birds are both nectivorous and frugivorous (i.e. “dual mutualists”) and (2) how avian translocations and conservation reestablished nectivory and frugivory networks. Results showed that all but one native bird species were dual mutualists. Pairwise species interaction frequencies were positively correlated between networks. However, overall levels of nectivory by each bird species were unrelated to levels of frugivory. Interaction specialization and species strength also did not differ between networks. The reintroduction of threatened and endangered bird species appeared to have restored both interaction networks, and the sequence of species recovery accelerated restorative changes. Overall results indicate that not only does the extinction of dual mutualists have a 2-fold, negative effect on mutualistic interactions with plants, they can also accelerate the recovery of ecosystem services following restoration efforts.     

Phenology of fleshy fruits in a wet sclerophyll forest in southeastern Australia: are birds an important influence?

Summary The hypothesis that seasonality in the production of fleshy fruits in temperate regions is the result of selection by avian seed dispersal agents or avian seed predators was tested in a temperate wet sclerophyll forest in south-eastern Australia. I determined whether ten common fleshy-fruited species produced fruits when potential avian seed dispersers were most abundant or avian seed predators were least abundant. The season of fruit production was not correlated with avian disperser abundance nor with seasonal changes in avian seed predators. Peak fruiting occurred during autumn but fruit-eating birds were equally abundant from spring through to autumn. Avian seed predators (parrots) did not show any seasonal variation in abundance. If dispersers are influencing fruiting seasonality in wet sclerophyll forest, it is likely that changes in social behaviour and feeding patterns of dispersers during autumn, rather than increases in abundance, will be an important influence on fruiting patterns in wet sclerophyll forest. However, environmental and life history factors may also influence phenology.     

Bat frugivory in two subtropical rain forests of Northern Argentina: Testing hypotheses of fruit selection in the Neotropics

Phyllostomid bats are prominent components of mammalian assemblages in the Neotropics. With many species specialized in frugivory, phyllostomids represent major partners of fleshy-fruited plants in the mutualism of seed dispersal. Here we present dietary data from two subtropical rainforests of Argentina, where fruit diversity is low and thus offer unique opportunities to test hypotheses of diet selection originally proposed for species-rich tropical assemblages. Particularly, we tested whether frugivorous phyllostomids exhibit pronounced dietary specialization in core plant taxa where fruit offer is greatly reduced as compared to tropical rainforests. We analyzed dietary overlap and niche breadth of subtropical frugivorous bats on the basis of >1000 dietary records plus >500 samples from a previous study in the region. We show that in the subtropics, frugivores from different genera remain faithful to their respective core plant taxa with few exceptions, rather than shifting toward alternative fruit resources available in the study sites. This supports predictions of specialization, which is confirmed to have a deep historical origin. The response of phyllostomid ensembles to restricted fruit diversity is at the level of species composition: absence of species for which preferred fruits do not occur in the sites. Taken together, these data lend strong support to hypotheses that explain coexistence of frugivorous phyllostomids on the basis of dietary specialization on core plant taxa with chiropterochorous fruits.     

Influence of type of avian frugivory on the fitness ofPistacia terebinthus L.

Summary The fruits ofPistacia terebinthus, a circum-Mediterranean tree/shrub, are consumed by an array of bird species that differ in feeding methods and in relative frequencies of visits to plants. In this study I document interindividual variation in the proportion of fruits consumed by three types of frugivores: legitimate dispersers, pulp-consumers and seed predators. The results show that the relative frequencies of each kind of frugivore notably influence the final reproductive output (absolute number of viable seeds dispersed) and in fact prevail over the effects of pre-dispersal factors acting on plant fitness. Those relative frequencies are not associated with any of the plant traits related to fitness, such as fruit crop size and the number of viable seeds produced, suggesting that the type of avian frugivory exerts a negligible, if not null, selective pressure on such plant attributes. Plant specialization to attract the most effective seed dispersers seems to be precluded, given the small scale at which the high variation in seed dispersal success takes place.     

Interspecific and annual variation in pre-dispersal seed predation by a granivorous bird in two East Asian hackberries, Celtis biondii and Celtis sinensis

Pre-dispersal seed predation by granivorous birds has potential to limit fruit removal and subsequent seed dispersal by legitimate avian seed dispersers in bird-dispersed plants, especially when the birds form flocks. We monitored pre-dispersal seed predation by the Japanese grosbeak, Eophona personata, of two bird-dispersed hackberry species (Cannabaceae), Celtis biondii (four trees) and Celtis sinensis (10 trees), for 3 years (2005, 2007 and 2008) in a fragmented forest in temperate Japan. Throughout the 3 years, predation was more intense on C. biondii, which, as a consequence, lost a larger part of its fruit crop. Grosbeaks preferred C. biondii seeds that had a comparatively lower energy content and lower hardness than C. sinensis, suggesting an association between seed hardness and selective foraging by grosbeaks. In C. biondii, intensive predation markedly reduced fruit duration and strongly limited fruit removal by seed dispersers, especially in 2007 and 2008. In C. sinensis, seed dispersers consumed fruits throughout the fruiting seasons in all 3 years. In C. biondii, variation in the timing of grosbeak migration among years was associated with annual variation in this bird's effects on fruit removal. Our results demonstrate that seed predation by flocks of granivorous birds can dramatically disrupt seed dispersal in fleshy-fruited plants and suggest the importance of understanding their flocking behaviour.     

Dietary specialization and fruit availability among frugivorous birds on Sulawesi

Over 20 months I collected data on the diet composition and fruit availability for 23 frugivorous bird species (pigeons, parrots, hornbills and passerines) in two lowland rain forests on Sulawesi, Indonesia. Using these data I determined species dependency on fruit, diet composition, diet breadth and the availability of their fruit resources. Fruit dependency ranged from 44 to 100% of diets across the assemblage, with significant inter-order differences reflecting shared evolutionary history within orders. By contrast, diet breadths did not differ between bird orders owing to high within-order variability, possibly as a result of species-specific foraging strategies. Overall, no significant relationship existed between fruit dependency and diet breadth, which indicated that they are not interchangeable measures of dietary specialization, but that each provided important information on dietary specialization, and enabled assessment of resource requirements and availability. The assemblage consumed the fruits of 120 species representing 40 plant families. Figs ( Ficus spp.) were of great importance across the assemblage and comprised 57% of all fruit-eating records. For hornbills, passerines and pigeons, figs accounted for substantial proportions of monthly feeding records (= 58%) and were consumed whatever the availability of other fruits. Figs were therefore a fundamental part of species' diets, possibly every month, and not merely important during times of relative scarcity of other fruit. Conservation of fig trees both inside and outside protected areas is likely to be important in maintaining healthy populations of figs and the frugivorous birds that depend on them.