Gibbons (Nomascus gabriellae) provide key seed dispersal for the Pacific walnut (Dracontomelon dao), in Asia's lowland tropical forest

Understanding the mutualisms between frugivores and plants is essential for developing successful forest management and conservation strategies, especially in tropical rainforests where the majority of plants are dispersed by animals. Gibbons are among the most effective seed dispersers in South East Asia's tropical forests, but are also one of the highly threatened arboreal mammals in the region. Here we studied the seed dispersal of the Pacific walnut (Dracontomelon dao), a canopy tree which produces fruit that are common in the diet of the endangered southern yellow-cheeked crested gibbon (Nomascus gabriellae). We found that gibbons were the most effective disperser for this species; they consumed approximately 45% of the fruit crop, which was four times more than that consumed by macaques – the only other legitimate disperser. Gibbons tracked the temporal (but not spatial) abundance of ripe fruits, indicating this fruit was a preferred species for the gibbon. Both gibbons and macaques dispersed the majority (>90%) of the seeds at least 20 m away from parent crowns, with mean dispersal distances by gibbons measuring 179.3 ± 98.0 m (range: 4–425 m). Seeds defecated by gibbons germinated quicker and at greater rates than seeds spat by macaques, or in undispersed fruits. Gibbon-dispersed seeds were also more likely to be removed by unknown seed predators or unknown secondary dispersers. Overall, gibbons play a key role in the regeneration of the Pacific walnut. Our findings have significant implications both for the management of the Pacific walnut tree dominating tropical rainforest as well as the reintroduction program of the Southern yellow-cheeked crested gibbon.     

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.     

Modified dispersal‐related traits in disjunct populations of bird‐dispersed Frangula alnus (Rhamnaceae): a result of its Quaternary distribution shifts?

Many European tree species survived Pleistocene glaciations in Mediterranean refugia and rapidly recolonized temperate Europe afterwards. Inter‐ and postglacial migration processes are assumed to have catalized evolutionary optimizations of dispersal‐related traits, but up to now empirical evidence is lacking in vertebrate‐dispersed plants. We investigated if south Iberian glacial relict and central European “colonizer” populations of the bird‐dispersed tree Frangula alnus have experienced differentiations of dispersal‐related traits which increase the mobility of northern populations. A comparison of lifetime reproductive strategy, disperser guilds, ripening phenology, and fruit design revealed considerable differences. Compared to south Iberian conspecifics, central European plants were considerably smaller and experienced a highly accelerated generation turnover. In south Iberian populations seed dispersal was carried out almost completely by resident birds which occurred in constant abundances throughout the ripening season. In contrast, central European seeds were dispersed by migrants whose abundances changed considerably during the ripening season. Several bird species were involved in both study areas but rendered different importance for seed dispersal. The fruit ripening pattern was highly asynchronous throughout the ripening season in south Iberia, while central European trees showed a complex ripening sequence which resulted in a significant correlation between fruit abundance and changing disperser availability. Central European fruits were smaller and showed a considerably smaller seed load than south Iberian fruits, thus presumably being more attractive for their small‐sized main dispersers (Sylvia warblers). Chemical analyses revealed significant differences in contents of water, glucose, fructose, proteins, ash. and phenolic compounds. The extensive differentiation of dispersal‐related traits in F. alnus suggests that even weak selective pressures by frugivores may induce evolutionary adjustments of dispersal traits over large time scales. We suggest that the differences we observe today evolved during the species' distribution shifts in the Quaternary.     

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.     

Do Meyer’s Parrots Poicephalus meyeri benefit pollination and seed dispersal of plants in the Okavango Delta,Botswana?

Recent studies of new world parrots repeatedly document, with few exceptions, that parrots are wasteful and destructive predispersal seed predators that are unlikely to contribute towards pollination and seed dispersal. Few detailed studies, however, have assessed the contribution of African parrots to forest ecology by quantifying the potential net benefit of seed and flower predation by parrots for most tree species in their diet. Due to the incidence of pollen on the heads of Meyer’s Parrots when feeding on Leguminosae flowers and the dispersal of viable seeds to the ground during seed predation, we compared destruction rates, when feeding on pods, fruits and flowers, with dispersal rates of viable seeds to the ground and frequency of head contact with reproductive apparatus to estimate net benefit from Meyer’s Parrot feeding activity. Meyer’s Parrots were not implicated in endo‐ or epizoochory, but they dropped uneaten fruit pulp and seeds to the ground during feeding bouts, thus providing ripe, undamaged seeds to secondary seed dispersers. This link with forest recruitment was weak, as all tree species utilized by Meyer’s Parrots either had more significant primary dispersal agents or were primarily wind‐dispersed. In most cases, the negative effect of seed predation outweighed any positive effects in terms of dispersal, whereby almost three times more seeds were consumed or destroyed than were dispersed to the ground. Significantly, only Sclerocarya birrea caffra recorded marginal net dispersal benefit from utilization by Meyer’s Parrots. Due to low relative resource abundance and high destruction rate, feeding activity on Berchemia discolor may be significant enough to influence its spatial distribution and abundance. Utilization of flowers of Kigelia africana and Adansonia digitata by parrots likely had a significant negative impact on pollination. Feeding on Acacia nigrescens flowers, however, was potentially advantageous to their pollination. We conclude that Poicephalus parrots are net consumers of ripe, undamaged seeds and flowers, thus having an overall negative impact on forest recruitment in subtropical Africa.     

The Role of Arboreal Seed Dispersal Groups on the Seed Rain of a Lowland Tropical Forest1

Most tropical plants produce fleshy fruits that are dispersed primarily by vertebrate frugivores. Behavioral disparities among vertebrate seed dispersers could influence patterns of seed distribution and thus forest structure. This study investigated the relative importance of arboreal seed dispersers and seed predators on the initial stage of forest organization–seed deposition. We asked the following questions: (1) To what degree do arboreal seed dispersers influence the species richness and abundance of the seed rain? and (2) Based on the plant species and strata of the forest for which they provide dispersal services, do arboreal seed dispersers represent similar or distinct functional groups? To answer these questions, seed rain was sampled for 12 months in the Dja Reserve, Cameroon. Seed traps representing five percent of the crown area were erected below the canopies of 90 trees belonging to nine focal tree species: 3 dispersed by monkeys, 3 dispersed by large frugivorous birds, and 3 wind‐dispersed species. Seeds disseminated by arboreal seed dispersers accounted for ca 12 percent of the seeds and 68 percent of the seed species identified in seed traps. Monkeys dispersed more than twice the number of seed species than large frugivorous birds, but birds dispersed more individual seeds. We identified two distinct functional dispersal groups, one composed of large frugivorous birds and one composed of monkeys, drop dispersers, and seed predators. These groups dispersed plants found in different canopy strata and exhibited low overlap in the seed species they disseminated. We conclude it is unlikely that seed dispersal services provided by monkeys could be compensated for by frugivorous birds in the event of their extirpation from Afrotropical forests.     

Non-native mammals are weak candidates to substitute ecological function of native avian seed dispersers in an island ecosystem

Although prominent examples exist of non-native species causing substantial ecological harm, many have neutral or positive effects, including filling surrogate roles once performed by extinct native organisms. We tested the ecological roles of two non-native mammals as seed dispersers or seed predators in Guåhan, which, due to invasive brown tree snakes (Boiga irregularis), is devoid of native seed dispersers–birds and bats. We conducted feeding trials with captive rats (Rattus spp.), which are present but uncommon due to predation by snakes, and pigs (Sus scrofa), which are abundant. We examined if and how they interacted with common forest fruits. We then compared how any gut-passed or animal-handled seeds germinated compared to seeds left in whole fruit or depulped seeds. Rats and pigs interacted with most of the fruits and seeds (>80%) that they were fed. Of those, most seeds were destroyed—78% for rats and 90% for pigs, across both native and non-native plant species. Compared to seeds germinating within whole fruits, rats improved germination of the seeds that they handled without ingesting, while pigs diminished the germination of seeds that they handled. The small percentage of seeds (approximately 1.5% for rats and 5% for pigs) that survived gut passage germinated in higher proportions than those in whole fruits. Percentages of seed survival to germination are lower than found in similar studies with native avian frugivores. Our results indicate that pigs and rats have mixed effects on seeds, but are not suitable surrogates for native seed dispersers.     

濒危树种闽楠种子和幼苗生态学研究

从1995至1998年,通过种子收集器布设、不同程度圈围的样方处理、实验室和野外发芽实验、相邻格子样方调查和土壤中种子的筛选等一系列方法,对福建罗卜岩闽楠种子质资源保护区内闽楠为优势的常绿阔叶林群落中闽楠种群的果实、种子库、种子散布、果实／种子捕良、种子萌发和幼苗存活等进行了研究。结果表明：1995至1996年果实量为116个／m^2,成熟高峰期在1996年1月中旬。1997年至1998年为103个/m^2,成熟高峰期在1997年12月中旬。1997年至1998年,绝大部分（90．1％）的果实直接从母树上掉落,低于9．5％的果实由鸟类传播。实验室的种子发芽率为93％,而在野外仅为12％。野外发芽率低多由种子霉烂引起。闽楠的种子寿命较短,其种子在次年4月初开始萌发,7月之后地面种子即丧夫活力。动物捕食不仅影响种子的存活,同时也影响幼苗的死亡率,未经圈围与样地中被取食的果实和幼苗数量有显著差异。在新形成的林窗中或远离母树均降低了果实的迁移和幼苗的死亡率。但各因素交互作用有所不同。密度相关效应发生在2－3个月的幼苗阶段,之后,密度高低对果实被取食和幼苗死亡没有显著的影响。这些结果表明,闽楠种子和幼苗经历较大的环境压力,即在高湿度生境中母树树冠下的种子不仅易受到土壤病原菌的感染,同样容易遭受动物所捕食,从而导致较低的野外种子发芽率和幼苗存活率。这是闽楠致濒的重要原因之一。     

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.     

Seed dispersal of Diospyros virginiana in the past and the present: Evidence for a generalist evolutionary strategy

Several North American trees are hypothesized to have lost their co‐evolved seed disperser during the late‐Pleistocene extinction and are therefore considered anachronistic. We tested this hypothesis for the American persimmon (Diospyros virginiana) by studying the effects of gut passage of proposed seed dispersers on seedling survival and growth, natural fruiting characteristics, and modern animal consumption patterns. We tested gut passage effects on persimmon seeds using three native living species, the raccoon (Procyon lotor), Virginia opossum (Didelphis virginiana), and coyote (Canis latrans), and two Pleistocene analogs; the Asian elephant (Elephas maximus) and alpaca (Vicugna pacos). Persimmon seeds excreted by raccoons, coyotes, and elephants survived gut transit. Gut passage did not affect sprouting success, but did tend to decrease time to sprout and increase seedling quality. Under field conditions, persimmon fruits were palatable on the parent tree and on the ground for an equal duration, but most fruits were consumed on the ground. Seven vertebrate species fed upon persimmon fruits, with the white‐tailed deer (Odocoileus virginianus)—a species not capable of dispersing persimmon seeds—comprising over 90% of detections. Conversely, potential living seed dispersers were rarely detected. Our results suggest the American persimmon evolved to attract a variety of seed dispersers and thus is not anachronistic. However, human‐induced changes in mammal communities could be affecting successful seed dispersal. We argue that changes in the relative abundance of mammals during the Anthropocene may be modifying seed dispersal patterns, leading to potential changes in forest community composition.     

Unexpected relationships and valuable mistakes: non‐myrmecochorous Prosopis dispersed by messy leafcutting ants in harvesting their seeds

Abstract Ants generally disperse seeds while feeding on fruits or structures attached to the seed. Seed dispersal as a by‐product of seed predation (dyszoochory) was recognized in specialized harvester ants, but not in ants predating seeds opportunistically. Leafcutting ants are the main herbivores in much of the Neotropics, and they have been reported to remove fruits and seeds, but their role as seed predators and dispersers has not been acknowledged. Prosopis flexuosa D.C. (Fabaceae, Mimosoideae) is the most abundant tree species in the central Monte Desert, Argentina, and it is likely to depend on secondary animal dispersal. Mammalian frugivores are usually considered its main dispersers, but the opportunity for dispersal may be small since the removal of fruits and seeds by seed predators is very intense. The objective of this study was to identify which ant species interact with P. flexuosa fruits and to evaluate their relative importance as seed predators and dispersers. In a field experiment, whole and segmented pods were offered and several ant species exploiting the fruits were identified. Additionally, all pod segments remaining around nests of the three ant species able to remove them (the leafcutters Acromyrmex lobicornis Emery and Acromyrmex striatus Roger, and Pheidole bergi Mayr) were examined during and after the P. flexuosa primary dispersal season. Up to 753 pod segments and 90 sound seeds were found accumulated in a circle of 1 m radius over nests of A. lobicornis, and even more in an examined trail. Acromyrmex striatus left a smaller proportion of sound seeds and P. bergi left a smaller number of pod segments. All tendencies were similar during shorter known periods of accumulation. Leafcutting ants are acting as important seed predators, and ‘by mistake’ may be dispersing a key non‐myrmecochorous tree. This is an unexplored path in the seed dispersal cycle of P. flexuosa that challenges the tendency to predict interactions based on classifications made with other goals.     

The role of avian ‘seed predators’ as seed dispersers

Seed dispersal is a central process in plant ecology with consequences for species composition and habitat structure. Some bird species are known to disperse the seeds they ingest, whereas others, termed ‘seed predators’, digest them and apparently play no part in dispersal, but it is not clear if these are discrete strategies or simply the ends of a continuum. We assessed dispersal effectiveness by combining analysis of faecal samples and bird density. The droppings of seed dispersers contained more entire seeds than those of typical seed predators, but over a quarter of the droppings of seed predators contained whole seeds. This effect was further magnified when bird density was taken into account, and was driven largely by one frequent interaction: the Chaffinch Fringilla coelebs, a typical seed predator and the most abundant bird species in the area and dispersed seeds of Leycesteria formosa, a non‐native plant with berry‐like fruits. These results suggest the existence of a continuum between seed predators and seed dispersers.     

Standard yet unusual mechanisms of long-distance dispersal: seed dispersal of Corymbia torelliana by bees

Mellitochory, seed dispersal by bees, has been implicated in long-distance dispersal of the tropical rain forest tree, Corymbia torelliana (Myrtaceae). We examined natural and introduced populations of C. torelliana for 4 years to determine the species of bees that disperse seeds, and the extent and distance of seed dispersal. The mechanism of seed dispersal by bees was also investigated, including fruit traits that promote dispersal, foraging behaviour of bees at fruits, and the fate of seeds. The fruit structure of C. torelliana , with seed presented in a resin reward, is a unique trait that promotes seed dispersal by bees and often results in long-distance dispersal. We discovered that a guild of four species of stingless bees, Trigona carbonaria, T. clypearis, T. sapiens , and T. hockingsi, dispersed seeds of C. torelliana in its natural range. More than half of the nests found within 250 m of fruiting trees had evidence of seed transport. Seeds were transported minimum distances of 20–220 m by bees. Approximately 88% of seeds were dispersed by gravity but almost all fruits retained one or two seeds embedded in resin for bee dispersal. Bee foraging for resin peaked immediately after fruit opening and corresponded to a peak of seed dispersal at the hive. There were strong correlations between numbers of seeds brought in and taken out of each hive by bees ( r =  0.753–0.992, P  < 0.05), and germination rates were 95 ± 5%. These results showed that bee-transported seeds were effectively dispersed outside of the hive soon after release from fruits. Seed dispersal by bees is a non-standard dispersal mechanism for C. torelliana, as most seeds are dispersed by gravity before bees can enter fruits. However, many C. torelliana seeds are dispersed by bees, since seeds are retained in almost all fruits, and all of these are dispersed by bees.     

Vertebrate Fruit Removal and Ant Seed Dispersal in the Neotropical Ginger Renealmia alpinia (Zingiberaceae)*

Plants frequently display fruit characteristics that support multiple seed‐dispersal syndromes. These ambiguous characteristics may reflect the fact that seed dispersal is usually a complex process involving multiple dispersers. This is the case for the Neotropical ginger Renealmia alpinia (Zingiberaceae). It was originally suggested that the aromatic fruits of R. alpinia located at the base of the plant are adapted for terrestrial mammal seed dispersal. However, the dark‐purple coloration of the fruits and bright orange aril surrounding the seeds suggest that birds may play a role in R. alpinia seed dispersal. At La Selva Biological Station, Costa Rica, we used camera traps to record vertebrate visits to infructescences of R. alpinia. Most visitors were toucans and aracaris (Ramphastidae). However fruits were also removed by terrestrial mammals (coatis and armadillos). In addition to vertebrate fruit removal, some of the fruits dehisce and the seeds that fall on the ground are dispersed by ants. Fruitfall traps showed that 77 percent of fruits are removed by vertebrates. However, 15 percent of fruits fall to the base of parent plants to be potentially dispersed by ants. Experiments using a laboratory ant colony showed that ants are effective seed dispersers of R. alpinia. Ant seed manipulation increased germination success and reduced time to germination. In conclusion, primary seed dispersal in the Neotropical ginger R. alpinia is mostly performed by birds, additionally ants are effective dispersers at short distances. Seed dispersal in R. alpinia is a complex process involving a diverse array of dispersal agents.     

Seed Dispersal in the Dark: Shedding Light on the Role of Fruit Bats in Africa

In spite of their recognized importance as seed dispersers in other parts of the tropics, seed dispersal by fruit bats has received scant research attention in Africa. To evaluate the role of African fruit bats in seed dispersal, we studied fruits and seeds below 480 bat feeding roosts in the East Usambara Mountains of Tanzania. We compared these findings to those reported in other African localities to place our results in a broader context. We found 49 plant species dispersed by bats: 28 species, 18 genera, and one family are novel reports of bat dispersal in Africa. Approximately 20 percent of the submontane tree flora of the East Usambaras is bat‐dispersed, including both widespread and endemic trees. African fruit bats are important seed dispersers at our study site because they move seeds of dozens of species tens or hundreds of meters, even seeds that are too large to ingest (greater than 5 mm in length). Fruit bats are likely important seed dispersers in other Afrotropical forests, as bats elsewhere in Africa are known to consume 20 genera and 16 species of plants reported here. Insights from studying remains under bat feeding roosts offer a simple method to further document and substantially increase our understanding of the role of African fruit bats in seed dispersal.     

Rare seed-predating mammals determine seed fate of <Emphasis Type="Italic">Canarium euphyllum</Emphasis>, a large-seeded tree species in a moist evergreen forest,Thailand

Natural seed deposition patterns and their effects on post-dispersal seed fate are critical to tropical tree recruitment. The major dispersal agents of the large-seeded tree Canarium euphyllum in Khao Yai National Park, Thailand, are large frugivorous birds such as hornbills, which generated spatially heterogeneous seed deposition patterns because they regurgitated seeds at perching trees and conspecific and heterospecific feeding trees. We investigated the fate of seeds dispersed in this manner using seed removal experiments and automatic camera trapping. Seeds placed experimentally around conspecific feeding trees had higher removal rates than seeds placed elsewhere. These effects were likely mediated by two seed-eating rodents, the Indochinese ground squirrel (Menetes berdmorei) and the giant long-tailed rat (Leopoldamys sabanus). Consequently, the spatial patterns generated by hornbills had consequences for post-dispersal seed fates, particularly whether or not the seeds were removed by rodents. Primary dispersal by hornbills does alter seed fate by altering the probability of rodent–seed interaction, but the ultimate impact of dispersal by hornbills will depend on how important rodent scatterhoarding is to seed germination and seedlings. Given that major seed dispersers of C. euphyllum are now absent or rare in degraded forests in tropical Asia, it is becoming increasingly important to understand the roles of scatterhoarding rodents in these altered habitats in this region.     

Cones structure and seed traits of four species of large‐seeded pines: Adaptation to animal‐mediated dispersal

Seed dispersal selection pressures may cause morphological differences in cone structure and seed traits of large‐seeded pine trees. We investigated the cone, seed, and scale traits of four species of animal‐dispersed pine trees to explore the adaptations of morphological structures to different dispersers. The four focal pines analyzed in this study were Chinese white pine (Pinus armandi), Korean pine (P. koraiensis), Siberian dwarf pine (P. pumila), and Dabieshan white pine (P. dabeshanensis). There are significant differences in the traits of the cones and seeds of these four animal‐dispersed pines. The scales of Korean pine and Siberian dwarf pine are somewhat opened after cone maturity, the seeds are closely combined with scales, and the seed coat and scales are thick. The cones of Chinese white pine and Dabieshan white pine are open after ripening, the seeds fall easily from the cones, and the seed coat and seed scales are relatively thin. The results showed that the cone structure of Chinese white pine is similar to that of Dabieshan white pine, whereas Korean pine and Siberian dwarf pine are significantly different from the other two pines and vary significantly from each other. This suggests that species with similar seed dispersal strategies exhibit similar morphological adaptions. Accordingly, we predicted three possible seed dispersal paradigms for animal‐dispersed pines: the first, as represented by Chinese white pine and Dabieshan white pine, relies upon small forest rodents for seed dispersal; the second, represented by Korean pine, relies primarily on birds and squirrels to disperse the seeds; and the third, represented by Siberian dwarf pine, relies primarily on birds for seed dispersal. Our study highlights the significance of animal seed dispersal in shaping cone morphology, and our predictions provide a theoretical framework for research investigating the coevolution of large‐seeded pines and their seed dispersers.     

Asymmetrical Dependence Between a Neotropical Mistletoe and its Avian Seed Disperser

The degree of interdependence among interacting species has major implications for our understanding of the coevolutionary process and biodiversity maintenance. However, the mutualism strength among fruiting plants and their seed dispersers remains poorly understood in tropical ecosystems. We evaluated simultaneously the effectiveness of the avian seed dispersers of the mistletoe Struthanthus flexicaulis (Loranthaceae) and the contribution of its fruits to their diets in a highland rocky savanna in southeastern Brazil. The mistletoe fruits are small lipid‐rich pseudoberries available throughout the year. Four passerine birds fed on fruits, but Elaenia cristata (Tyrannidae) was the most effective disperser, responsible for more than 96 percent of the dispersed seeds. This bird swallowed fruits whole, expelling and depositing undamaged seeds by regurgitation and bill wiping on perches. From 646 dispersed seeds, 56 percent were deposited on safe sites, thin live twigs of 38 susceptible host species. Elaenia cristata were predominantly frugivorous, feeding on typically ornithocoric fruits of at least 12 species, but also on arthropods. Although fruits represented 75 percent of the feeding bouts along the year, S. flexicaulis fruits represented only 34 percent of the E. cristata diet. Our results highlight the asymmetrical nature of this mutualistic interaction, with the mistletoe life cycle locally linked to one highly effective seed disperser that is more weakly dependent on mistletoes fruits as a food source. We suggest that merging the seed dispersal effectiveness framework with diet assessment of seed dispersers is needed to clarify the asymmetries in mutualistic pairwise interactions involving plants and their animal partners.     

How intraspecific variation in seed‐dispersing animals matters for plants

Seed dispersal by animals is a complex phenomenon, characterized by multiple mechanisms and variable outcomes. Most researchers approach this complexity by analysing context‐dependency in seed dispersal and investigating extrinsic factors that might influence interactions between plants and seed dispersers. Intrinsic traits of seed dispersers provide an alternative way of making sense of the enormous variation in seed fates. I review causes of intraspecific variability in frugivorous and granivorous animals, discuss their effects on seed dispersal, and outline likely consequences for plant populations and communities. Sources of individual variation in seed‐dispersing animals include sexual dimorphism, changes associated with growth and ageing, individual specialization, and animal personalities. Sexual dimorphism of seed‐dispersing animals influences seed fate through diverse mechanisms that range from effects caused by sex‐specific differences in body size, to influences of male versus female cognitive functions. These differences affect the type of seed treatment (e.g. dispersal versus predation), the number of dispersed seeds, distance of seed dispersal, and likelihood that seeds are left in favourable sites for seeds or seedlings. The best‐documented consequences of individual differences associated with growth and ageing involve quantity of dispersed seeds and the quality of seed treatment in the mouth and gut. Individual specialization on different resources affects the number of dispersed plant species, and therefore the connectivity and architecture of seed‐dispersal networks. Animal personalities might play an important role in shaping interactions between plants and dispersers of their seeds, yet their potential in this regard remains overlooked. In general, intraspecific variation in seed‐dispersing animals often influences plants through effects of these individual differences on the movement ecology of the dispersers. Two conditions are necessary for individual variation to exert a strong influence on seed dispersal. First, the individual differences in traits should translate into differences in crucial characteristics of seed dispersal. Second, individual variation is more likely to be important when the proportions of particular types of individuals fluctuate strongly in a population or vary across space; when proportions are static, it is less likely that intraspecific differences will be responsible for changes in the dynamics and outcomes of plant–animal interactions. In conclusion, focusing on variation among foraging animals rather than on species averages might bring new, mechanistic insights to the phenomenon of seed dispersal. While this shift in perspective is unlikely to replace the traditional approach (based on the assumption that all important variation occurs among species), it provides a complementary alternative to decipher the enormous variation observed in animal‐mediated seed dispersal.     

Complementary roles of two resilient neotropical mammalian seed dispersers

Capuchin monkeys (Cebus spp. and Sapajus spp.) and coatis (Nasua spp.) coexist in most neotropical forests, including small forest remnants. Both capuchins and coatis eat fruit and disperse seeds, but little is known about whether their roles in seed dispersal are redundant or complementary. We compiled 49 studies from the literature on feeding by capuchins and/or coatis, of which 19 were comprehensive enough for our analyses. We determined the relative importance of fruit eating to each species and compared their diets. Additionally, we analysed the structure of three fruit–frugivore networks built with both animal groups and the fruits they eat and evaluated whether fruit traits influenced the network topology. Fruits represented the largest part of capuchin and coati diets, even though coatis have been known for their opportunistic and generalist diets. Capuchins and coatis also exhibited similar general diet parameters (niche breadth and trophic diversity). The three networks exhibited high connectance values and variable niche overlap. A Multiple Correspondence Analysis, failed to detect any trait or trait combination related to food use. In conclusion, capuchins and coatis both have generalist diets; they feed on many different species of fruits and exhibit important complementarity as seed dispersers. Both are likely to be particularly important seed dispersers in disturbed and fragmented forests.