Reliability of macaques as seed dispersers

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

Persistence of the effect of frugivore identity on post‐dispersal seed fate: consequences for the assessment of functional redundancy

Large frugivores play an important role as seed dispersers and their extinction may affect plant regeneration. The consequences of such extinctions depend on the likelihood of other species being functionally redundant and on how post‐dispersal events are affected. We assess the functional redundancy of two seed dispersers of the Atlantic Forest, the muriqui (Brachyteles arachnoides) and the tapir (Tapirus terrestris) through the comparison of their seed dispersal quality, taking into account post‐dispersal events. We compare tapirs and muriquis for: (1) the dung beetle community associated with their feces; (2) the seed burial probability and burial depth by dung beetles; and (3) the seed mortality due to predators or other causes according to burial depth. We determine how seed burial affects seed dispersal effectiveness (SDE) and compare the dispersal quality of four plant species dispersed by these frugivores. Muriqui feces attract 16‐fold more dung beetles per gram of fecal matter and seeds experience 10.5‐fold more burial than seeds in tapir feces. In both feces types, seed mortality due to predation decreases with burial depth but seed mortality due to other causes increases. Total seed mortality differ within plant species according to the primary disperser. Therefore, the effect of seed burial on SDE varies according to the plant species, burial depth, and primary disperser. As tapirs and muriquis differently affect the seed fate, they are not functionally redundant. Since the effect of the primary disperser persists into the post‐dispersal events, we should consider the cascading effects of these processes when assessing functional redundancy.     

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.     

Negative density dependence of seed dispersal and seedling recruitment in a Neotropical palm

Negative density dependence (NDD) of recruitment is pervasive in tropical tree species. We tested the hypotheses that seed dispersal is NDD, due to intraspecific competition for dispersers, and that this contributes to NDD of recruitment. We compared dispersal in the palm Attalea butyracea across a wide range of population density on Barro Colorado Island in Panama and assessed its consequences for seed distributions. We found that frugivore visitation, seed removal and dispersal distance all declined with population density of A. butyracea, demonstrating NDD of seed dispersal due to competition for dispersers. Furthermore, as population density increased, the distances of seeds from the nearest adult decreased, conspecific seed crowding increased and seedling recruitment success decreased, all patterns expected under poorer dispersal. Unexpectedly, however, our analyses showed that NDD of dispersal did not contribute substantially to these changes in the quality of the seed distribution; patterns with population density were dominated by effects due solely to increasing adult and seed density.     

Forest cover,hunting pressure,and fruit availability influence seed dispersal in a forest‐savanna mosaic in the Congo Basin

Forest fragmentation, reduced forest cover, and hunting pressure are the main threats affecting animal‐mediated seed dispersal. However, their combined effects on seed dispersal rates have been simultaneously investigated only rarely, and never in Africa. We aimed to disentangle the effects of forest cover, hunting pressure, frugivore abundance, and fruit availability at the local and landscape scales on the seed dispersal rates of Staudtia kamerunensis (Myristicaceae). To estimate the percentages of seed dispersal failure (undispersed seeds), we quantitated fruit remains below fruiting trees distributed across five contrasting sites in a semi‐natural forest‐savanna mosaic in the Democratic Republic of Congo. We used statistical analyses accounting for spatial autocorrelation and found that forest cover in the surrounding landscape, hunting level, the associated abundance of dispersers, and fruit availability all had significant effects on the percentage of seed dispersal failure. The combination of high fruit availability and reduced abundance of seed dispersers could accelerate seed disperser satiation, causing the seed dispersal system to be saturated. Our study highlights how two major factors associated with anthropogenic activities, forest cover and hunting, affect seed dispersal by animals. These findings could have far‐reaching implications for our understanding of tree‐frugivore interactions and the conservation of tropical communities.     

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.     

Primary Seed Dispersal by Red Howler Monkeys and the Effect of Defecation Patterns on the Fate of Dispersed Seeds1

The effectiveness of a seed disperser depends on the quantity and quality of dispersal. The quality of dispersal depends in large part on factors that affect the post–dispersal fate of seeds, and yet this aspect of dispersal quality is rarely assessed. In the particular case of seed dispersal through endozoochory, the defecation pattern produced has the potential of affecting the fate of dispersed seeds and consequently, dispersal quality and effectiveness. In this study, I assessed the effects of dung presence and dung/seed densities on seed predation by rodents and secondary dispersal by dung beetles. In particular, I compared seed fates in clumped defecation patterns, as those produced by howler monkeys, with seed fates in scattered defecation patterns, as those produced by other frugivores. I also determined the prevalence of red howler monkeys (Alouatta seniculus) as seed dispersers at the plant community level in Central Amazonia by determining the number of species they dispersed in a 25–month period. I found that dung presence and amount affected rodent and dung beetle behavior. Seed predation rates were higher when dung was present, and when it was in higher densities. The same number of seeds was buried by dung beedes, in dumped versus scattered defecation patterns, but more seeds were buried when they were inside large dung–piles versus small piles. Seed density had no effect on rodent or dung beetle behavior. Results indicate that caution should be taken when categorizing an animal as a high or low quality seed disperser before carefully examining the factors that affect the fate of dispersed seeds. Red howler monkeys dispersed the seeds of 137 species during the study period, which is the highest yet reported number for an Alouatta species, and should thus be considered highly prevalent seed dispersers at the plant community level in Central Amazonian terra firme rain forests.     

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.     

Spatial Pattern of Pindó Palm (Syagrus romanzoffiana) Recruitment in Argentinian Atlantic Forest: The Importance of Tapir and Effects of Defaunation

Hunting pressure, fragmentation and deforestation have caused global declines in animal abundance, and the consequences for plant communities are poorly understood. Many large‐seeded plants, for instance, depend on large and endangered vertebrates for seed dispersal. In some Semi‐deciduous Atlantic Forests, endangered tapirs (Tapirus terrestris) are major dispersers of pindó palms (Syagrus romanzoffiana). Here, we compare recruitment patterns of pindó palms between protected and disturbed (defaunated) Atlantic Forest areas in Argentina and evaluate the potential consequences of the lack of the main disperser for pindó palm regeneration. We analyzed the number and spatial pattern of pindó adults, offspring, and tapir dung piles within ten plots established in an area spanning tapir latrines inside Iguazú National Park and in a fragmented forest area outside the park where tapir is locally extinct. In both areas, we evaluated recruitment levels beneath 24 adult palms in circular plots centered on adult stems. We found lower pindó palm recruitment outside the park where offspring tended to be aggregated around adult palms. In contrast, in Iguazú National Park offspring were spatially associated with tapir dung‐piles, in which most offspring were registered. Recruitment under adults was higher outside the park suggesting a lower rate of seed removal in disturbed areas. Our results show that tapir dispersal promotes higher recruitment levels of pindó offspring and shapes their spatial pattern, breaking the spatial association with adult (presumably maternal) palms. These results are useful for predicting the impact of local tapir extinction on this palm.     

Tamarins and Dung Beetles: An Efficient Diplochorous Dispersal System in the Peruvian Amazonia

Dung beetles fulfill several key functions in ecosystems but their role as secondary seed dispersers is probably one of the most complex ones. Various factors, such as seed characteristics, dispersal pattern induced by the primary disperser, season, and habitat, can affect the seed–beetle interaction. Particularly little is known about the fate of seeds primarily dispersed in small feces. The aim of this study was to investigate the effects of these factors on the dung beetle community (species composition, number and size of individuals) and its consequences on burial occurrence and depth of seeds primarily dispersed by two tamarin species. We captured dung beetles in a Peruvian rain forest with 299 dung‐baited pitfall traps to characterize the dung beetle community. Seed burial occurrence and depth were assessed by marking in situ 551 dispersed seeds in feces placed in cages. Among these seeds, 22.5 percent were buried by dung beetles after 2 d. We observed a significant effect of the amount of dung, season, time of deposition, and habitat on the number of individuals and species of dung beetles, as well as on seed burial occurrence and depth, while the tamarin species significantly influenced only the number and the size of dung beetles. This seed dispersal loop is particularly important for forest regeneration: small to large seeds dispersed by tamarins in secondary forest can be buried by dung beetles. These seeds can thus benefit from a better protection against predation and a more suitable microenvironment for germination, potentially enhancing seedling recruitment.     

Intraspecific variation in fruit–frugivore interactions: effects of fruiting neighborhood and consequences for seed dispersal

The extent of specialization/generalization continuum in fruit–frugivore interactions at the individual level remains poorly explored. Here, we investigated the interactions between the Neotropical treelet Miconia irwinii (Melastomataceae) and its avian seed dispersers in Brazilian campo rupestre. We built an individual-based network to derive plant degree of interaction specialization regarding disperser species. Then, we explored how intraspecific variation in interaction niche breadth relates to fruit availability on individual plants in varying densities of fruiting conspecific neighbors, and how these factors affect the quantity of viable seeds dispersed. We predicted broader interaction niche breadths for individuals with larger fruit crops in denser fruiting neighborhoods. The downscaled network included nine bird species and 15 plants, which varied nearly five-fold in their degree of interaction specialization. We found positive effects of crop size on visitation and fruit removal rates, but not on degree of interaction specialization. Conversely, we found that an increase in the density of conspecific fruiting neighbors both increased visitation rate and reduced plant degree of interaction specialization. We suggest that tracking fruit-rich patches by avian frugivore species is the main driver of density-dependent intraspecific variation in plants’ interaction niche breadth. Our study shed some light on the overlooked fitness consequences of intraspecific variation in interaction niches by showing that individuals along the specialization/generalization continuum may have their seed dispersed with similar effectiveness. Our study exemplifies how individual-based networks linking plants to frugivore species that differ in their seed dispersal effectiveness can advance our understanding of intraspecific variation in the outcomes of fruit–frugivore interactions.

     

Dispersión Primaria de Semillas por Primates y Dispersión Secundaria por Escarabajos Coprófagos en Tikal,Guatemala1

We linked primary dispersal by spider monkeys (Ateles geoffroyi) and howler monkeys (Alouatta pigra) to post‐dispersal seed fate by studying the effects of dung type and defecation pattern on secondary seed dispersal by dung beetles. First, we described the defecation patterns for both primate species. Howler monkeys generally defecated in groups (88% of observed defecations), with each individual producing on average 31 g of dung, resulting in a large area of the forest floor (31 m²) covered by large amounts of dung (clumped spatial pattern). Spider monkeys generally (96% of observed defecations) defecated individually, each individual producing an average of 11 g of dung, resulting in a small area of the forest floor (2 m²) covered by small amounts of dung (scattered spatial pattern). Secondly, we captured dung beetles using as bait the dung of both primate species, to detect differences in the assemblages of these secondary seed dispersers attracted to the dung of both primates. More individual dung beetles, but not more species, were attracted to howler monkey dung than to spider monkey dung. Finally, we assessed experimentally (using plastic beads as seed mimics) how dung type (Ateles vs. Alouatta) and defecation pattern (scattered vs. clumped) affect secondary seed dispersal by dung beetles. We found that post‐dispersal seed fate was affected by dung type, with more seeds being buried when present in howler monkey dung, than in spider monkey dung, but was not affected by defecation pattern. It is important to consider post‐dispersal processes, such as secondary seed dispersal by dung beetles, when comparing species of primary dispersers.     

Changes of effective gene dispersal distances by pollen and seeds across successive life stages in a tropical tree

Pollen and seed dispersal are the two key processes in which plant genes move in space, mostly mediated by animal dispersal vectors in tropical forests. Due to the movement patterns of pollinators and seed dispersers and subsequent complex spatial patterns in the mortality of offspring, we have little knowledge of how pollinators and seed dispersers affect effective gene dispersal distances across successive recruitment stages. Using six highly polymorphic microsatellite loci and parentage analyses, we quantified pollen dispersal, seed dispersal, and effective paternal and maternal gene dispersal distances from pollen‐ and seed‐donors to offspring across four recruitment stages within a population of the monoecious tropical tree Prunus africana in western Kenya. In general, pollen‐dispersal and paternal gene dispersal distances were much longer than seed‐dispersal and maternal gene dispersal distances, with the long‐distance within‐population gene dispersal in P. africana being mostly mediated by pollinators. Seed dispersal, paternal and maternal gene dispersal distances increased significantly across recruitment stages, suggesting strong density‐ and distance‐dependent mortality near the parent trees. Pollen dispersal distances also varied significantly, but inconsistently across recruitment stages. The mean dispersal distance was initially much (23‐fold) farther for pollen than for seeds, yet the pollen‐to‐seed dispersal distance ratio diminished by an order of magnitude at later stages as maternal gene dispersal distances disproportionately increased. Our study elucidates the relative changes in the contribution of the two processes, pollen and seed dispersal, to effective gene dispersal across recruitment. Overall, complex sequential processes during recruitment contribute to the genetic make‐up of tree populations. This highlights the importance of a multistage perspective for a comprehensive understanding of the impact of animal‐mediated pollen and seed dispersal on small‐scale spatial genetic patterns of long‐lived tree species.     

Context‐dependent fruit–frugivore interactions: partner identities and spatio‐temporal variations

Fruit–frugivore interactions are crucial for the dynamics and regeneration of most forested ecosystems. Still, we lack an understanding of the potential variation in the sign and strength of such interactions in relation to variations in the spatial and temporal ecological context. Here, we evaluated spatial (three sites) and temporal (two fruiting seasons) local variation in the sign (seed predation versus dispersal) and strength (frequency and quantity) of the interactions among six frugivorous mammals and a community of Mediterranean fleshy‐fruited shrubs. We examined mammal faecal samples and quantified frequency of seed occurrence, number of seeds per faecal sample, seed species diversity and quality of seed treatment (i.e. percentage of undamaged seeds). The frequency of seed occurrence and number of seeds per faecal sample strongly varied among dispersers, sites, seasons and fruit species. For instance, fox Vulpes vulpes faeces showed between 6 and 40 times more seeds than wild boar Sus scrofa faeces in seasons or sites in which Rubus and Juniperus seeds were dominant. However, in seasons or sites dominated by Corema seeds, wild boar faeces contained up to seven times more seeds than fox faeces. Mammalian carnivores (fox and badger, Meles meles) treated seeds gently, acting mostly as dispersers, whereas deer (Cervus elaphus and Dama dama) acted mainly as seed predators. Interestingly, rabbit Oryctolagus cuniculus acted as either mostly seed disperser or seed predator depending on the plant species. Our results indicated that the sign of fruit–frugivore interactions depended mainly on the identity of the partners. For a particular fruit–frugivore pair, however, our surrogate of interaction strength largely varied with the spatio‐temporal context (year and habitat), leading to a low specificity across the seed–frugivore network. The high spatio‐temporal variability of seed dispersal (in quantity, quality and seed diversity) by different frugivores would confer resilience against unpredictable environmental conditions, such as those typical of Mediterranean ecosystems.     

Lek Breeding and Territorial Aggression in White-eared Kob

Territorial aggression in a lek breeding population of white-eared kob (Kobus kob leucotis) was investigated in the Boma National Park region, southern Sudan. The frequency of aggression on leks was positively related to the number of females present, but generally declined over the course of the breeding season. Males fought most strenuously for central territories that were preferred by females. Males with females in their territories were more frequently engaged in fights than unaccompanied males. Such fights often induced females to leave their original partners, especially when large groups of females were involved. Territorial aggression led to damaging injuries in several instances, and mortality of breeding age males was disproportionately high. These results suggest that the intensity of aggression exhibited by territorial males was scaled to potential reproductive benefits.     

Dung Beetle Communities and Seed Dispersal in Primary Forest and Disturbed Land in Amazonia1

Seeds from tropical fruiting trees ingested and defecated on the soil surface by primary dispersers (such as primates) are vulnerable to destruction from rodents, insects, and fungi. Burial by dung beedes as an incidental result of their feeding and nesting activities often provides these seeds with refugia from attack. To examine the effect of habitat disturbance on the dung beetle communities involved in this process, I surveyed dung beedes at three sites in the Amazon basin, in die states of Parí, Amazonas, and Rondónia, Brazil. Through principal component analysis on measurements of size and behavioral characters of beedes, I determined the relative quality of beetle species as seed dispersers (dispersal defined as horizontal or vertical movement of seeds) and ranked them into seed dispersal guilds. I used correspondence analysis to examine in what habitats (primary forest or varying degrees of disturbed habitat) these guilds were abundant. Most guilds decreased with increasing habitat disturbance, but one guild made up of large nocturnal burrowers (primarily Dichotomius) became more abundant with increasing disturbance (up to the level of highest disturbance surveyed), at which point all dung beede species became scarce. Clear–cuts had lower species richness, lower abundance, and lower biomass dian forested areas. These results imply that clear–cutting is detrimental to all seed dispersal dynamics in tropical rain forests, but that some levels of disturbance allow enough disperser activity to preserve this ecosystem function.     

Effects of forest fragmentation on the oak–rodent mutualism

Land‐use changes are expected to affect plant–disperser conditional mutualisms through changes in animal behavior. We analyzed the oak–rodent conditional mutualism in Mediterranean fragmented forests at two climatically different locations. We quantified fragmentation effects on seed dispersal effectiveness and assessed if such effects were due to changes in habitat structure and intraspecific competition for acorns in fragmented areas. Fragmentation decreased cover from predators within mouse territories as well as intraspecific competition for acorns. This resulted in lower dispersal effectiveness in small forest fragments. Globally, habitat structure was the main driver in mouse foraging decisions. In small fragments, low shelter availability precluded mouse movements, leading to short mobilization distances and low caching rates. However, as the proportion of cover from predators increased, mice were able to modulate their foraging decisions depending on intraspecific competition for acorns, resulting in higher dispersal quality. In addition to fragmentation effects, delayed breeding in the southern locality caused lower number of rodents during the dispersal season, which reduced acorn mobilization rates. Our study shows that seed dispersal patterns in managed systems can be analyzed as the result of management effects on key environmental factors in dispersers’ foraging decisions.     

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.     

Impact of seed abundance on seed processing and dispersal by the red ant Myrmica rubra

1. Myrmecochory sensu stricto is an ant–plant mutualism in which non‐granivorous ants disperse plant diaspores after feeding on their nutrient‐rich seed appendage, the elaiosome. Phenological traits associated with the diaspore can influence the behaviour of ants and thus their ultimate efficiency as seed dispersers. 2. This study investigated how a contrasting availability of seeds (20 vs. 200 seeds) from the diplochorous Chelidonium majus (Papaveraceae, Linnaeus) plant species influences the behaviour of Myrmica rubra (Formicidae, Linnaeus) ants, from the retrieval of seeds until their dispersal outside the ant nest. 3. Regardless of seed abundance, the ants collected the first diaspores at similar rates. Then, seed retrieval sped up over time for large seed sources until satiation took place with only one‐third of the tested colonies wholly depleting abundant seed sources. 4. No active recruitment by trail‐laying ants was triggered, even to an abundant seed source 5. In both conditions of seed abundance, the majority of the diaspores retrieved inside the nest were discarded with the elaiosome removed and were dispersed at similar distances from the nest. 6. The paper concludes with a discussion of how the quantity of seeds released by a plant with a dual mode of dispersal can potentially influence the behaviour of ant dispersers and hence the dispersal efficiency derived from myrmecochory.     

Effects of zoochory on the spatial genetic structure of plant populations

Spatial genetic structure (SGS) of plants results from the nonrandom distribution of related individuals. SGS provides information on gene flow and spatial patterns of genetic diversity within populations. Seed dispersal creates the spatial template for plant distribution. Thus, in zoochorous plants, dispersal mode and disperser behaviour might have a strong impact on SGS. However, many studies only report the taxonomic group of seed dispersers, without further details. The recent increase in studies on SGS provides the opportunity to review findings and test for the influence of dispersal mode, taxonomic affiliation of dispersers and their behaviour. We compared the proportions of studies with SGS among groups and tested for differences in strength of SGS using Sp statistics. The presence of SGS differed among taxonomic groups, with reduced presence in plants dispersed by birds. Strength of SGS was instead significantly influenced by the behaviour of seed dispersal vectors, with higher SGS in plant species dispersed by animals with behavioural traits that result in short seed dispersal distances. We observed high variance in the strength of SGS in plants dispersed by animals that actively or passively accumulate seeds. Additionally, we found SGS was also affected by pollination and marker type used. Our study highlights the importance of vector behaviour on SGS even in the presence of variance created by other factors. Thus, more detailed information on the behaviour of seed dispersers would contribute to better understand which factors shape the spatial scale of gene flow in animal‐dispersed plant species.