The role of avian frugivores in germination of seeds of fleshy-fruited invasive alien plants

Many highly invasive plant species have fleshy fruits which are eaten by native frugivorous animals. These frugivores play an important role in long-distance seed dispersal, and may also affect germination success. The aim of this study was to determine whether generalist frugivores enhance or decrease seed germination of invasive alien species through pulp removal or seed coat abrasion, besides serving as dispersal agents. Fruits of four fleshy-fruited invasive alien plant species, namely Solanum mauritianum, Cinnamomum camphora, Lantana camara and Psidium guajava, were fed to three generalist avian frugivorous species, which have been observed feeding on these fruits in the wild. Seed retention time was recorded as this affects dispersal distance and the duration that seeds are exposed to the effects of the gut. Seeds removed from excreta, seeds from manually de-pulped fruit, and whole fruit were planted in soil trays housed in a greenhouse. Daily germination counts were done. Seed retention times differed significantly between bird species for all fruits, except those of C. camphora. However, all frugivores had a similar effect on the germination success of seeds of S. mauritianum, L. camara and P. guajava, showing that gut retention time was not important. Germination of seeds from manually de-pulped fruits did not differ from that of ingested seeds of all plant species, suggesting that seed coat abrasion was also not important. Pulp removal resulted in significantly higher germination rates, both in the two species with larger, multi-seeded fruit (S. mauritianum and P. guajava), and in the two species having single-seeded fruit with waxy exocarps (C. camphora and L. camara). Pulp removal also resulted in significantly earlier germination of L. camara and P. guajava seeds. Therefore, frugivores not only accelerate dispersal, but also greatly enhance seed germination of all fleshy-fruited invasive alien species in this study.     

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.     

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.     

Revealing secondary seed removers: results from camera trapping

This paper reports the results of the first study on secondary seed removal of seeds dispersed by Sykes’ monkeys (Cercopithecus albogularis) using camera traps in Africa. Patterns of primary seed dispersal are often superimposed by secondary conveyance, emphasising the need to study these secondary processes carefully. As the agents and mechanisms of seed dispersal are often concealed, being carried out by cryptic or nocturnal animals in dense vegetation, camera trapping was deemed a viable means to investigate secondary removal of seeds disseminated by C. albogularis in the Western Soutpansberg, South Africa. Camera traps were established at preferred feeding trees of the focal Sykes’ monkey group to identify animal species that remove seeds and fruits spat and dropped to the forest floor and seed removal observations were carried out. This method proved to be effective in identifying seed removers and also allowed to get indications about the quantities of seeds removed. Ten animal species were recorded visiting the trees, of which eight were observed removing seeds and fruits. Overall seed and fruit removal rates were high, indicating that the foraging behaviour of C. albogularis attracts many terrestrial frugivores.     

Impact of avian frugivores on dispersal and recruitment of the invasive <Emphasis Type="Italic">Prunus serotina</Emphasis> in an agricultural landscape

Although seed dispersal is considered to be a key process determining the spatial structure and spread of non-native plant populations, few studies have explicitly addressed the link between dispersal vector behaviour, seed distribution and seedling recruitment to gain insight into the process of exotic species invasion within a fragmented landscape context. The present study analyses the relationship between avian frugivory and spatial patterns of seed deposition and seedling recruitment for an expanding population of the invasive Prunus serotina in a hedgerow network landscape in Flanders, Belgium. We quantified fruit production, observed frugivores, and determined the spatial distribution of bird droppings and P. serotina seedlings. A relatively diverse assemblage of frugivores visited P. serotina seed trees, with Columba palumbus and Turdus merula being by far the most important dispersers. Landscape structure strongly affected dispersal vector behaviour and the spatial distribution of perching birds, droppings and seedlings. Frugivorous birds non-randomly dispersed seeds to perching sites and an association between perching birds, seed deposition and seedling recruitment was found. Results indicate that landscape structure contributes to non-random seed deposition of P. serotina by common local frugivores. Cutting the larger seed trees is proposed as the most feasible measure to slow down the invasion rate.     

Tree dispersal strategies in the littoral forest of Sainte Luce (SE-Madagascar)

Zoochory is the most common mode of seed dispersal for the majority of plant species in the tropics. Based on the assumption of tight plant-animal interactions several hypotheses have been developed to investigate the origin of life history traits of plant diaspores and their dispersers, such as species-specific co-evolution, the low/high investment model (low investment in single fruits but massive fruiting to attract many different frugivores versus high investment in single fruits and fruit production for extended periods to provide food for few frugivores), and the evolution of syndromes which represent plant adaptations to disperser groups (e.g. birds, mammals, mixed). To test these hypotheses the dispersal strategies of 34 tree species were determined in the littoral forest of Sainte Luce (SE-Madagascar) with the help of fruit traps and tree watches. The impact of fruit consumers on the seeds was determined based on detailed behavioral observations. Phenological, morphological and biochemical fruit traits from tree species were measured to look for co-variation with different types of dispersal. No indication for species-specific co-evolution could be found nor any support for the low/high investment model. However dispersal syndromes could be distinguished as diaspores dispersed by birds, mammals or both groups (mixed) differ in the size of their fruits and seeds, fruit shape, and seed number, but not in biochemical traits. Five large-seeded tree species seem to depend critically on the largest lemur, Eulemur fulvus collaris, for seed dispersal. However, this does not represent a case of tight species-specific co-evolution. Rather it seems to be the consequence of the extinction of the larger frugivorous birds and lemurs which might also have fed on these large fruits. Nevertheless these interactions are of crucial importance to conserve the integrity of the forest.     

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.     

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.     

Weak interactions between avian and insect frugivores: the case of Pistacia terebinthus L. (Anacardiaceae)

Vertebrate frugivores often feed on fruits upon or within which insects also feed, yet little information exists on the potential magnitude of interactions between these consumers. The Mediterranean shrub Pistacia terebinthus, the birds that consume its fruits, and the wasps that feed upon its seeds are examined in this study. P. terebinthus produces a highly variable fraction of final-sized red fruits that never become mature (green-colored). Red fruits can be immature, parthenocarpic, aborted, or attacked by wasps, and their pulp is much less nutritious than that of mature fruits. A total of 20 bird species consumed the fruits in the study area. Legitimate dispersers accounted for 39% of the total fruit removal, while pulp eaters and seed predators accounted for the remainder. Birds strongly preferred the mature fruits (only 4% of the fruits consumed were red). The incidence of wasps in the seeds ranged from 0 to 42% of the crop in 1989 and from 0 to 24% in 1990. The influence of avian and insect frugivore guilds on each other appears to be quite low because of the narrow overlap in resource utilization by birds and wasps, and an overall low intensity of wasp seed predation. From an evolutionary perspective, the possible ability of wasps to preclude fruit maturation appears not to be attributable to the present interaction with avian frugivores.     

Exploring the interaction of avian frugivory and plant spatial heterogeneity and its effect on seed dispersal kernels using a simulation model

Seed dispersal by avian frugivores is one of the key processes influencing plant spatial patterns, but may fail if there is disruption of plant–frugivore mutualisms, such as decline in abundance of dispersers, fragmentation of habitat, or isolation of individual trees. We used simulation model experiments to examine the interaction between frugivore density and behaviour and the spatial arrangement of fruiting plants and its effect on seed dispersal kernels. We focussed on two New Zealand canopy tree species that produce large fruits and are dispersed predominantly by one avian frugivore (Hemiphaga novaeseelandiae). Although the mean seed dispersal distance decreased when trees became more aggregated, there were more frugivore flights between tree clusters, consequently stretching the tails of the dispersal kernels. Conversely, when trees were less aggregated in the landscape, mean dispersal distances increased because seeds were deposited over larger areas, but the kernels had shorter tails. While there were no statistically meaningful changes in kernel parameters when frugivore density changed, decreases in density did cause a proportional reduction in the total number of dispersed seeds. However, birds were forced to move further when fruit availability and fruit ripening were low. Sensitivity analysis showed that dispersal kernels were primarily influenced by the model parameters relating to disperser behaviour, especially those determining attractiveness based on distance to candidate fruiting trees. Our results suggest that the spatial arrangement of plants plays an important role in seed dispersal processes – although tree aggregation curbed the mean seed dispersal distance, it was accompanied by occasional long distance events, and tree dispersion caused an increase in mean dispersal distance, both potentially increasing the probability of seeds finding suitable habitats for germination and growth. Even though low frugivore densities did not cause dispersal failure, there were negative effects on the quantity of seed dispersal because fewer seeds were dispersed.     

Role of small rodents in the seed dispersal process: Microcavia australis consuming Prosopis flexuosa fruits

Understanding the functional role of animal species in seed dispersal is central to determining how biotic interactions could be affected by anthropogenic drivers. In the Monte Desert, mammals play different functional roles in Prosopis flexuosa seed dispersal, acting as opportunistic frugivores (endozoochorous medium‐sized and large mammals) or seed hoarders (some small sigmodontine rodents). Our objective was assessing the functional role of Microcavia australis, a small hystricognathi rodent, in the fruit removal and seed deposition stages of P. flexuosa seed dispersal, compared to sympatric sigmodontine rodents. In situ, we quantified fruit removal by small rodents during non‐fruiting and fruiting periods, and determined the distance seeds were transported, particularly by M. australis. In laboratory experiments, we analysed how M. australis stores seeds (through scatter‐ or larder‐hoarding) and how many seeds are left in caches as living seeds, relative to previous data on sigmodontine rodents. To conduct field studies, we established sampling stations under randomly chosen P. flexuosa trees at the Ñacuñán Man and Biosphere Reserve. We analysed fruit removal by small rodents and seed dispersal distance by M. australis using camera traps focused on P. flexuosa fruits covered with wire screen, which only allowed entry of small animals. In laboratory trials, we provided animals with a known number of fruits and assessed seed conditions after removal. Small rodents removed 75.7% of fruit supplied during the non‐fruiting period and 53.2% during the fruiting period. Microcavia australis and Graomys griseoflavus were the main fruit removers. Microcavia australis transported seeds to a mean distance of 462 cm and cached seeds mainly in scatter‐hoards, similarly as Eligmodontia typus. All transported seeds were left in fruit segments or covered only by the endocarp, never as predated seeds. Microcavia australis disperses P. flexuosa seeds by carrying fruits away from a source to consume them and then by scatter‐hoarding fruits and seeds.     

Composition and clumping of seeds deposited by frugivorous birds varies between forest microsites

More frequent deposition of seeds by frugivores beneath plants in fruit could impose spatial limits to the distribution of plants dispersed by animals and contribute to species coexistence. Also, differences in diet and use of microhabitats by seed dispersers could promote spatial variation in the combination of seed species deposited. We investigated patterns of seed deposition of Miconia fosteri and Miconia serrulata (Melastomataceae) by birds in the Amazon. The goal was to determine how distribution and abundance of fruiting plants, both con‐ and hetero‐specifics, affect the spatial variability in clumping and composition of multi‐specific seed deposition. We established two 9‐ha plots in undisturbed terra‐firme understory in the Ecuadorian Amazon. Seed rain was sampled with seed traps located in four microsites: below plants of the focal species, below Anthurium eminens (Araceae), and in randomly selected microsites. We examined seed deposition in these microsites in relation to habitat, fruiting neighborhood (fruit abundance, and distance to and density of plants of the target species), and crop size of M. fosteri or M. serrulata to determine if microsites differed in abundance and species composition of seeds. Seed traps below plants in fruit received more seeds than did randomly located traps. Seeds of the target species were, moreover, more commonly deposited below con‐ rather than hetero‐specific plants. Seed aggregation below fruiting plants increased in forest neighborhoods where the abundance of fruits and the combination of fruiting plant species contributed to the arrival of seeds. Microsites differed notably in the combination of seeds deposited by frugivores, and differences were less pronounced among microsites that received seeds of M. fosteri and M. serrulata than among all microsites where at least some seed species were deposited by birds. We demonstrate that two closely related, ecologically similar species possess many similarities in their patterns of seed deposition and in the factors that affect those patterns. The combination of seed species deposited below foci of dispersal depended on the fruiting plant species, and the spatial patterns of seed deposition varied with the location of the microsite and the combination of co‐dispersed species in the neighborhood. Similar species that share the same dispersers were confronted with different combinations of seeds depending on the microsite where they arrived, which could promote forest heterogeneity in the combination of plant species.     

Patterns of frugivory in the columnar cactus Pilosocereus leucocephalus

In the frugivory networks of many arid and semi‐arid Mesoamerican ecosystems, columnar cacti act as keystone species that produce fruits with a high content of water and nutrients attractive to numerous vertebrates. The aim of this investigation was to assess the fruit removal patterns of two guilds of frugivores on the fruits of the woolly torch Pilosocereus leucocephalus. We assessed fruit pulp removal in two ways: by estimating the consumption of seeds given the amount of pulp removed per visit and by estimating the percentage of pulp removal over time. We put exclosures on unripe, intact fruits to keep frugivores from removing material. Once ripe, we removed the exclosures and tracked animal visitation of 69 fruits using camera traps. We obtained a total of 2,162 hr of footage (14:47 hours of them with effective pulp removal). The highest number of visitors is that of diurnal species (n = 12, all birds) versus only four nocturnal (three bats, one rodent). The most effective species in pulp removal are birds. Bats play a modest role in frugivory of this cactus. The significance of this work is twofold: (a) birds and bats consume the fruit pulp of this cactus and likely disperse its seeds, and (b) although bats rank high in pulp removal effectiveness, birds as a guild far outweigh their importance in this system, as they are not only more frequent but also remove more pulp and seeds. Both groups are known to be important in cacti seed dispersal, and our findings are essential in understanding the population dynamics of the woolly torch and in elucidating its seed dispersal ecology.     

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

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

Complementary seed dispersal by three avian frugivores in a fragmented Afromontane forest

Questions: To what extent does species‐specific variation in gut passage time (GPT), habitat use and mobility of three key avian frugivores synergistically affect the distribution of Xymalos monospora seeds within and among isolated forest fragments? Location: Three fragments of a severely fragmented cloud forest, Taita Hills, southeast Kenya. Methods: We experimentally determined GPTs of X. monospora seeds and recorded movements and habitat use by Turdus helleri, Andropadus milanjensis and Tauraco hartlaubi through radiotelemetry, and combined these data to generate species‐specific seed dispersal patterns. Results: Differences in mobility and habitat use among the three frugivores caused significant complementarity in seed dispersal, despite the fact that gut transit times were highly comparable. While the most sedentary and forest‐dependent species mainly led to short‐distance dispersal away from parent trees, two more mobile species dispersed seeds further away from the source trees, both within indigenous forest patches and towards exotic plantations and isolated fruiting trees in the landscape matrix. A. milanjensis inhabiting a very small forest fragment spent significantly more time in the landscape matrix than conspecifics residing in the two larger fragments. Conclusions: By varying distances over which seeds are carried away from parent trees and the habitat types in which they are ultimately deposited, avian frugivores affect the spatial distribution of seeds and early plant recruits in a distinct and complementary manner. Because landscape properties are expected to lead to different constraints on avian mobility for habitat specialists and for generalists, ecosystem processes such as avian seed dispersal are shaped by complex interactions between disperser behaviour and the environment.     

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.     

Wahlberg’s epauletted fruit bat (<Emphasis Type="Italic">Epomophorus wahlbergi</Emphasis>) as a potential dispersal agent for fleshy-fruited invasive alien plants: effects of handling behaviour on seed germination

The spread of invasive alien plants into natural habitats is of growing global concern. Several studies have investigated the role that avian frugivores play in the dispersal of these seeds and their effects on germination success. Fruit bats have however received little attention as important dispersal agents of invasive alien plants, despite their recognized role as long distance dispersal agents of various native flora. We investigated whether Wahlberg’s epauletted fruit bats, Epomophorus wahlbergi, would positively influence the germination of seeds of invasive alien plants. These fruit bats were fed fruits of four invasive alien plant species—Psidium guajava, Melia azedarach, Eriobotrya japonica, and Morus alba. Epomophorus wahlbergi were able to process more fruit per gram body mass than birds have been observed to do. Spat and de-pulped control seeds had similar germination success and germinated at approximately the same time for most species. While seeds retained in whole fruit had significantly less germination success than spat seeds for all species, except M. azedarach, they mostly germinated at approximately the same time. Epomophorus wahlbergi can swallow small seeds (< 2 mm), while seeds larger than this are generally spat out. Large fruit are usually carried away to feeding roosts where seeds are dropped, thereby dispersing seeds and fruits which are too large for some bird species to ingest. Epomophorus wahlbergi should not be underestimated as dispersers of these invasive alien plants as they consume proportionally large amounts (0.62 ± 0.09 to 0.99 ± 0.11 g.g⁻¹ body mass) of fruit, except for M. azedarach, and positively affect their seed germination rates.     

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.     

Avian fruit consumption and seed dispersal in a temperate Australian woodland

Abstract The effectiveness of avian fruit consumers as seed dispersers of fleshy‐fruited plants was studied in a temperate woodland community. As a consequence of the short and overlapping fruiting phenologies of fleshy‐fruited plant species in temperate regions of Australia, there are very few avian species that are true specialist frugivores. The relative importance of bird species as fruit consumers was investigated, and how their foraging activities, movements and gut treatment of seeds affected dispersal of viable seeds away from the parent plant was examined. Fruit consumption and consumer seed dispersal capacity were assessed in this study through faecal analyses and by testing the viability of seeds that had passed through the gut of avian consumers. Behavioural observations enabled us to determine the consumption rates of, and quantities of fruit consumed by, various bird species and the amount of time spent feeding. Silvereyes (Zosterops lateralis) were the dominant fruit consumers in the community, although 19 bird species were either observed consuming fruit or provided faecal samples that contained fruit. Silvereyes had a high local abundance at the site and more than 90% of silvereyes’faecal samples contained the seeds of fruiting plants (n = 409). Large numbers of fruit were consumed per visit by silvereyes, particularly for Rhagodia parabolica (fragrant saltbush). Silvereyes consumed an average of four R. parabolica fruit per 5 s and up to a maximum 40 fruit per visit. Viability was high for seeds recovered from silvereyes’faeces (R. parabolica, 94.4% viable; Hymenanthera dentata, 100% viable). However, the number of seeds per faecal sample was high for R. parabolica, which may result in density‐dependent seed mortality. Gut passage rate for silvereyes fed R. parabolica fruit in captivity was 31.5 ± 1.9 min. Silvereyes remained at fruiting plants for very short periods (average 50‐60 s) and in most cases moved away from the parent plant, primarily toward canopy trees. Given the short visit duration of silvereyes, individuals would have left the parent plant well before seeds passed through the gut. Rhagodia parabolica fruit was consumed by a large number of bird species in the community, including species often thought of as exclusively insectivorous or nectarivorous. These species are likely to disperse viable R. parabolica seeds into microhabitats different from those visited by silvereyes.     

The effect of frugivory on postdispersal seed removal and germination in the pantropical forest tree Antiaris toxicaria Leschenault

The quality of seed treatment by frugivores has an effect on seed removal after dispersal, seed germination and tree recruitment. We provide information on postdispersal seed removal, germination and subsequent recruitment in tropical forest tree species Antiaris toxicaria in Ghana. We tested whether postdispersal seed removal and germination rates were differentially affected by the following seed treatments: seeds that were spat out by monkeys with all fruit pulp removed and spitting seeds with fruit pulp partially removed as observed in some birds and bats. We used seeds of intact ripened fruits as control. Frugivore seed treatment and distance from bole affected seed removal patterns, whereas intact seeds were significantly removed from all seed stations. The germination success was greater for seeds that were spat out by monkeys and poor for seeds with fruit pulp partially removed and intact fruits. More recruits were recorded at the edge of the adult A. toxicaria canopy radius. There was weak relationship (r² = 0.042) between the number of recruits and distance away from the adult tree. Results suggest that the subsequent recruitment in tropical forest tree species may be enhanced by some frugivore fruit‐handling behaviour where fruit pulp is removed from the seeds without destroying the seeds.