Effective nut dispersal by magpies (<Emphasis Type="Italic">Pica pica</Emphasis> L.) in a Mediterranean agroecosystem

Scatter-hoarding animals such as corvids play a crucial role in the dispersal of nut-producing tree species. This interaction is well known for some corvids, but remains elusive for other species such as the magpie (Pica pica), an abundant corvid in agroecosystems and open landscapes of the Palearctic region. In addition, the establishment of the individual dispersed seeds—a prerequisite for determining seed-dispersal effectiveness—has never before been documented for the interaction between corvids and nut-producing trees. We analyzed walnut dispersal by magpies in an agroecosystem in southern Spain. We used several complementary approaches, including video recording nut removal from feeders, measuring dispersal distance using radio tracking (with radio transmitters placed inside nuts), and monitoring the fate of dispersed nuts to the time of seedling emergence. Magpies were shown to be highly active nut dispersers. The dispersal distance averaged 39.6 ± 4.5 m and ranged from 4.1 to 158.5 m. Some 90% of the removed walnuts were cached later, and most of these (98%) were buried in the soil or hidden under plant material. By the time of seedling emergence, ca. 33% of nuts remained at the caching location. Finally, 12% of the cached nuts germinated and 4% yielded an emerged seedling, facilitating the transition to the next regeneration stage. The results demonstrate for the first time that magpies can be an effective scatter-hoarding disperser of a nut-producing tree species, suggesting that this bird species may play a key role in the regeneration and expansion of broadleaf forests in Eurasia.     

Nut predation and dispersal of Harland Tanoak Lithocarpus harlandii by scatter-hoarding rodents

Plants that use the propagule to co-opt animals as dispersal agents must balance the costs of seed predation with the benefits of dispersal. Successful post-dispersal germination is a key metric that reflects these costs and benefits. By tracking individual nuts with coded tin-tags over 3 years (2000–2003), this study quantified nut predation and dispersal of harland tanoak (Lithocarpus harlandii) by seed-caching rodents in a subtropical evergreen broadleaved forest in the Duiangyan Region of Sichuan Province, Southwest China. We found that tanoak seedlings established from rodent-generated caches in the primary stands over a 12-month post-dispersal period. Our results indicate that seed-caching rodents are effective dispersers of tanoak nuts, but dispersal effectiveness varies among years and stands, probably due to mast seeding of harland tanoak or community-level seed availability according to the predator satiation hypothesis. Some nut traits in tanoak species, e.g. large seed size, hard nut husk, lower tannin and mast seeding, are important characteristics for seed dispersal by scatter-hoarding rodents, compared with oak species with higher tannin content.     

Spatial patterns of dispersal, seed predation and germination during colonization of abandoned grassland by Quercus petraea and Corylus avellana

Key processes during recruitment of Quercus petraea and Corylus avellana were investigated in abandoned calcareous grasslands and adjacent scrub using the following methods: (1) observation of hoarding animals during the main period of ripening of acorns and hazel nuts, (2) exposition of seeds on the soil surface and in 5–6 cm depth to test differences in predation and germination, and (3) mapping of seedlings in the grasslands. European jays (Garrulus glandarius) and mice were the main disperser of seeds. Jays preferred acorns, whereas the rodents were less selective, but probably more important for dispersal of nuts. The maximum dispersal distance was about 10–20 m for mice and was estimated several hundred metres for jays. Mice collected hoards of several seeds in about 2 cm depth in the soil, whereas jays stored single seeds. Seed predation and probably hoarding by mice were highest under scrub and in unmown grassland, while jays preferred mown sites for hoarding. However, hiding of seeds in the soil reduced losses in all sites. Predation of nuts was slightly less intensive than that of acorns. Seeds of Corylus were more sensitive to desiccation than Quercus, but in both species germination was lower for seeds exposed on the soil surface and in drier sites. Quercus and Corylus were the most abundant woody species in the fallow grasslands, probably due to the effective multi-staged dispersal by jays and mice, whereas wind-dispersed and fleshy-fruited species were less common; the latter restricted to margins of adjacent scrubland. The study provides several examples for discordance in suitability of patches for seeds and seedlings due to different habitat requirements of successive developmental stages. This result emphasizes the need for studies in the multiple stages during recruitment of vertebrate-dispersed plants.     

动物对松属植物种子的传播作用研究进展

松属植物约110种,根据种子传播方式可分为风传播松和动物传播松。风传播松占绝大多数,种子多具有适应风力的翅。动物传播松大约23种,都具有大、可食用、无翅或短翅的种子,无法借助风力传播。动物传播松的分布生境多为贫瘠的山地,而且多位于高海拔地区。目前已知9种松树的动物传播种类,其余14种可推测为动物传播。动物传播者包括鸦科鸟类和啮齿类动物,动物将获得的种子分散贮藏,未被重取的种子可能萌发,完成传播。动物传播是定向传播,微生境多适合种子萌发。啮齿类的传播距离可达数10 m,而鸦科鸟类的传播距离可达数公里。动物传播的松树会出现树丛和多树干现象,一般由同一贮点内贮藏的多粒种子萌发造成的。动物贮藏的种子大部分被重取,称传播后取食。一些具有大种子的风传播松在种子落地后,啮齿类和鸟类会再次埋藏而形成二次传播,可看做是一个单独的传播类型,即风-动物传播松。动物传播者与依赖传播松树之间可看作是互利共生关系。     

Dispersal of sweet pignut hickory in a year of low fruit production,and the influence of predation by a curculionid beetle

Summary The rate at which fallen hickory nuts are removed from beneath the parent tree, and the effect on this rate of the seed predatorConotrachelus affinis, was studied in an oak-hickory forest in southeastern Michigan, USA, during a year in which few nuts were produced. The trees responded toConotrachelus, which destroyed half the nut crop, by aborting inviable nuts during the summer. The seed dispersers, mostly gray squirrels, removed fallen nuts rapidly, showing the ability to distinguish viable nuts and remove them preferentially. The number of nuts removed in a week varies directly with the number available, and removal rate increases when many viable nuts are falling. The death of most seeds before dispersal, and the squirrels' efficiency at foraging on nuts and recovering them after burial, imply that successful hickory reproduction takes place only in years of heavy nut production.     

Acorn dispersal by the blue jay (Cyanocitta cristata)

Summary Blue jays transported and cached 133,000 acorns from a stand of Quercus palustris trees in Blacksburg, Virginia, representing 54% of the total mast crop. A further 20% (49,000) of the mast crop was eaten by jays at the collecting site. A large proportion of the nuts remaining beneath the collecting trees was parasitized by curculionid larvae. The number of nuts transported per caching trip ranged from 1–5 with a mean of 2.2. Mean distance between seed trees and caches was 1.1 km (range: 100 m–1.9 km). Jays appeared to choose species with small- to medium-sized nuts (Quercus palustris, Q. phellos, Q. velutina, Fagus grandifolia) and avoided the larger nuts of Q. borealis and Q. alba.Nuts were cached singly within a few meters of each other and were always covered with debris. Covering may improve germination and early growth by protecting the nut and radicle from desiccation. The vegetation structure of most suburban caching sites was analogous to open, disturbed environments in more natural landscapes. The presence of numerous Quercus seedlings in jay caching sites and the tendency for jays to cache nuts in environments conducive to germination and early growth indicate that blue jays facilitate colonization of members of the Fagaceae.     

Intraspecific directed deterrence by the mustard oil bomb in a desert plant

Plant secondary metabolites (SMs) acting as defensive chemicals in reproductive organs such as fruit tissues play roles in both mutualistic and antagonistic interactions between plants and seed dispersers/predators. The directed-deterrence hypothesis states that SMs in ripe fruits deter seed predators but have little or no effect on seed dispersers. Indeed, studies have demonstrated that birds are able to cope with fruit SMs whereas rodents are deterred by them. However, this mechanism was only demonstrated at the class level, i.e., between birds and mammals, based on differences in the vanilloid receptors. Here we present experimental and behavioral data demonstrating the use of the broad-range, class-independent "mustard oil bomb" mechanism in Ochradenus baccatus fruits to force a behavioral change at an ecological timescale, converting rodents from seed predators to seed dispersers. This is achieved by a unique compartmentalization of the mustard oil bomb, causing activation of the system only upon seed and pulp coconsumption, encouraging seed dispersal via seed spitting by rodents. Our findings demonstrate the power of SMs to shift the animal-plant relationship from predation to mutualism and provide support for the directed-deterrence hypothesis at the intraspecific level, in addition to the interspecific level.     

啮齿动物对米心水青冈种子命运的影响

有性生殖(实生更新)和无性生殖(萌生更新)是植物繁殖的两种关键方式。自然界中,个别物种同时具备两种更新方式,米心水青冈(Fagus engleriana Seem.)就是典型的物种。已有研究表明萌生更新在米心水青冈种群生活史中普遍存在,可使其占据原有生态位(生态位占据假说),但对米心水青冈实生更新的研究却未见报道。为了探索啮齿动物对米心水青冈实生更新的作用机制,笔者在神农架米心水青冈林中通过坚果摆放实验,研究啮齿动物对米心水青冈坚果传播机制的影响。结果显示,鼠类对坚果的原地捕食率高达81.22%;坚果被扩散率低,仅18.56%;坚果在原地平均留存时间为(5.25±5.95)d;坚果扩散距离为(2.63±1.60)m。研究发现,米心水青冈坚果被啮齿动物捕食率极高,贮藏率极低,且坚果被传播距离在同属植物中较小,不利于该物种更新繁殖。萌生更新在很大程度上弥补了米心水青冈实生更新繁殖方式的不足。本研究结果有助于对水青冈属植物繁殖策略的理解。     

Effects of seed size of wind-dispersed pines (Pinus) on secondary seed dispersal and the caching behavior of rodents

Seeds and nuts dispersed by scatter-hoarding animals are relatively large compared to propagules dispersed by other means. Possible selective forces in the evolution of large seed size include the selectivity of foraging animals and the ways that food-storing animals treat seeds and nuts after harvest. Treatment by rodents, primarily yellow pine chipmunks ( Tamias amoenus ), of four species of pine seeds that vary in size was studied in the Carson Range of western Nevada. The pines, lodgepole pine ( Pinus contorta , 8.7 mg), ponderosa pine ( P. ponderosa , 55 mg), Jeffrey pine ( P. jeffreyi , 157 mg), and sugar pine ( P. lambertiana , 213 mg), produce winged seeds that are initially wind-dispersed but are gathered by rodents and cached in the soil. Radioactive scandium-46 was used to follow the fates of seeds of all fours species placed around three source trees during autumn 1998 to 2000. Rodents gathered the seeds of all four species, but they took fewer of the lodgepole pine seeds and only six lodgepole seed caches (n=2106 total caches) were found during the three years. Among the other three species, number of seeds per cache decreased with increasing seed mass. However, the product of number of seeds per cache and seed mass was similar for all species. Sugar pine seeds were cached slightly deeper than ponderosa and Jeffrey pine seeds. For the species examined, seed size appeared to have had little effect on several other attributes, including mean dispersal distance, substrate choice, and microhabitat choice. Large size decreases wind dispersibility of pine seeds, but secondary dispersal by scatter-hoarding rodents compensates for poor wind dispersal so that total dispersibility of large-seeded pines is not compromised.     

Functional traits determine formation of mutualism and predation interactions in seed-rodent dispersal system of a subtropical forest

Network structure in plant-animal systems has been widely investigated but the roles of functional traits of plants and animals in formation of mutualism and predation interactions and community structure are still not fully understood. In this study, we quantitatively assessed interaction strength of mutualism and predation between 5 tree species and 7 rodent species by using semi-natural enclosures in a subtropical forest in southwest China. Seeds with high handling-time and nutrition traits (for both rat and mouse species) or high tannin trait (for mouse species) show high mutualism but low predation with rodents; while seeds with low handling-time and low nutrition traits show high predation but low mutualism with rodents. Large-sized rat species are more linked to seeds with high handling-time and high nutrition traits, while small-sized mouse species are more connected with seeds with low handling-time, low nutrition value and high tannin traits. Anti-predation seed traits tend to increase chance of mutualism instead of reducing predation by rodents, suggesting formation of mutualism may be connected with that of predation. Our study demonstrates that seed and animal traits play significant roles in the formation of mutualism and predation and network structure of the seed-rodent dispersal system.     

Potential for ants and vertebrate predators to shape seed-dispersal dynamics of the invasive thistles <Emphasis Type="Italic">Cirsium arvense</Emphasis> and <Emphasis Type="Italic">Carduus nutans</Emphasis> in their introduced range (North America)

The population dynamics of invasive plants are influenced by positive and negative associations formed with members of the fauna present in the introduced range. For example, mutualistic associations formed with pollinators or seed dispersers may facilitate invasion, but reduced fitness from attack by native herbivores can also suppress it. Since population expansion depends on effective seed dispersal, interactions with seed dispersers and predators in a plant species introduced range may be of particular importance. We explored the relative contributions of potential seed dispersers (ants) and vertebrate predators (rodents and birds) to seed removal of two diplochorous (i.e., wind- and ant-dispersed), invasive thistles, Cirsium arvense and Carduus nutans, in Colorado, USA. We also conducted behavior trials to explore the potential of different ant species to disperse seeds, and we quantified which potential ant dispersers were prevalent at our study locations. Both ants and vertebrate predators removed significant amounts of C. arvense and C. nutans seed, with the relative proportion of seed removed by each guild varying by location. The behavior trials revealed clear seed preferences among three ant species as well as differences in the foragers’ abilities to move seeds. In addition, two ant species that acted as potential dispersal agents were dominant at the study locations. Since local conditions in part determined whether dispersers or predators removed more seed, it is possible that some thistle populations benefit from a net dispersal effect, while others suffer proportionally more predation. Additionally, because the effectiveness of potential ant dispersers is taxon-specific, changes in ant community composition could affect the seed-dispersal dynamics of these thistles. Until now, most studies describing dispersal dynamics in C. arvense and C. nutans have focused on primary dispersal by wind or pre-dispersal seed predation by insects. Our findings suggest that animal-mediated dispersal and post-dispersal seed predation deserve further consideration.     

Sowing forests: a synthesis of seed dispersal and predation by agoutis and their influence on plant communities

Granivorous rodents have been traditionally regarded as antagonistic seed predators. Agoutis (Dasyprocta spp.), however, have also been recognized as mutualistic dispersers of plants because of their role as scatter-hoarders of seeds, especially for large-seeded species. A closer look shows that such definitions are too simplistic for these Neotropical animals because agoutis can influence plant communities not only through seed dispersal of large seeds but also through predation of small seeds and seedlings, evidencing their dual role. Herein, we summarize the literature on plant–agouti interactions, decompose agouti seed dispersal into its quantitative and qualitative components, and discuss how environmental factors and plant traits determine whether these interactions result in mutualisms or antagonisms. We also look at the role of agoutis in a community context, assessing their effectiveness as substitutes for extinct megafaunal frugivores and comparing their ecological functions to those of other extant dispersers of large seeds. We also discuss how our conclusions can be extended to the single other genus in the Dasyproctidae family (Myoprocta). Finally, we examine agoutis’ contribution to carbon stocks and summarize current conservation threats and efforts. We recorded 164 interactions between agoutis and plants, which were widespread across the plant phylogeny, confirming that agoutis are generalist frugivores. Seed mass was a main factor determining seed hoarding probability of plant species and agoutis were found to disperse larger seeds than other large-bodied frugivores. Agoutis positively contributed to carbon storage by preying upon seeds of plants with lower carbon biomass and by dispersing species with higher biomass. This synthesis of plant–agouti interactions shows that ecological services provided by agoutis to plant populations and communities go beyond seed dispersal and predation, and we identify still unanswered questions. We hope to emphasise the importance of agoutis in Neotropical forests.     

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.     

Experimental field test of spatial variation in rodent predation of nuts relative to distance and seed density

The spatial context in which seed predation occurs may modify the spatial structure of recruitment generated by seed dispersal. The Janzen–Connell (J-C) model predicts that granivores will exert greater pressure on the parent plant or at those sites where the density of dispersed seeds is higher. We have investigated how the probability of post-dispersal survival of Juglans australis varies with nut density across a hierarchy of spatial scales. We experimentally evaluated the survival of 3,120 nuts at three spatial scales: meso-scale (≤1.5 ha), as forest sites with two densities of fruiting J. australis individuals; intermediate scale (<0.2 ha), as individual trees with two experimental crop sizes; small scale (<0.1 m²), as microsites with two factors (number of nuts and distance from source). Nut removal coincided with seed predation, a condition that allowed us to test the density-dependent seed predation hypothesis. We found that the probability of nut survival was greater at forest sites with higher J. australis density. Nut survival was not affected by nut density in the seed shadow of individual specimens: at sites where J. australis density was greater, the proportion of surviving nuts did not differ between microsites located at different distances from the parent plant, but it was greater at microsites with greater initial nut density. Nut survival depended on the scale at which rodents responded to nut density, being negatively density dependent at the meso-scale and spatially random at intermediate and small scales. At the meso-scale, excess nut supply increased the probability of nut survival, which is in agreement with a model of granivore satiation near the seed source. Rodent satiation at the meso-scale may favour maintenance of sites with high J. australis density, where individual trees may have greater probabilities of passing their genes onto the next stage of the dispersal cycle.     

Look before leaping: foraging selectivity of capuchin monkeys on acacia trees in Costa Rica

Acacia trees in Costa Rica have an obligate mutualism with three species of Pseudomyrmex ants, which vigorously defend their host tree from insect and mammalian herbivores. Depending on the size and species of ant colony, individual acacia trees may be differentially protected. For animals able to discern between weakly and highly aggressive ant colonies, costs of ant stings from less active colonies might be offset by nutritional value acquired from feeding on acacia fruit or ant larvae in swollen thorns. We examined foraging selectivity of capuchin monkeys on acacia trees in Santa Rosa National Park, Costa Rica. We measured four characteristics of the acacia trees from which capuchins fed and of acacias immediately adjacent to those in which the monkeys fed: diameter at breast height (DBH), accessibility, species of closest tree and ant species present. We found that capuchins prefer to forage in acacias that are large and accessible. We also made two measurements of ant colony activity on each tree, one before and one after disturbing the ant colony. We found that the three species of mutualistic ants differ in baseline activity levels and that mutualistic ants are more active than non-mutualistic ant species found in acacia trees. We also found that capuchins foraged more frequently in trees colonized by non-mutualistic ants, but the explanatory value (r ²) of this model was low. Furthermore, monkeys did not discriminate between acacias on the basis of baseline ant activity or the ant colony’s response to disturbance. We conclude that these monkeys select acacia trees in which to forage based on characteristics of the trees rather than the ants. In addition, our study suggests that white-faced capuchins act as predators on the acacia ants but they probably benefit the dispersal and reproductive success of acacia trees. Capuchins may in fact function as an additional mutualistic partner for acacia trees via seed dispersal, but they must overcome the ants’ defense of the trees to do so.     

Geographic variation in nut size of<Emphasis Type="Italic"> Castanopsis</Emphasis> species in Japan

The environmental conditions affecting the geographic variation in nut size of the Castanopsis species in Japan were examined. The mother trees having a relatively large nut tended to dominate in the geographically marginal area of the distribution range in Japan, i.e. the Japan Sea, Kanto, and Ryukyu areas, whereas in the central area, i.e. the Pacific area, mother trees having a relatively small nut occurred. We attempted an adaptive explanation, proposing that mother trees with large nuts have higher inclusive fitness than those with small nuts in unfavorable habitats. The Japan Sea area features large amounts of snow fall and low temperatures. The Ryukyu area is insular and is characterized by high temperatures. Within the Pacific area, mother trees with relatively large nuts were abundant at both higher and lower parts of the altitudinal range. We infer that environmental conditions in marginal areas, in terms of both horizontal and vertical distribution, favor mother trees having large nuts. However, in the marginal Kanto area, the cause of the absence of mother trees having relatively small nuts was independent of the environmental conditions. The geographic variation in nut size of Castanopsis species is not always explained by environmental factors.     

Seed Dispersers,Seed Predators,and Browsers Act Synergistically as Biotic Filters in a Mosaic Landscape

In this study, we analize the functional influence of animals on the plants they interact with in a mediterranean mountain. We hypothesise that seed dispersers, seed predators, and browsers can act as biotic filters for plant communities. We analyse the combined effects of mutualistic (seed dispersal) and antagonistic (seed predation, herbivory) animal interactions in a mosaic landscape of Mediterranean mountains, basing our results on observational and experimental field. Most of the dispersed seeds came from tree species, whereas the population of saplings was composed predominantly of zoochorous shrub species. Seed predators preferentially consumed seeds from tree species, whereas seeds from the dominant fleshy-fruited shrubs had a higher probability of escaping these predators. The same pattern was repeated among the different landscape units by browsers, since they browsed selectively and far more intensely on tree-species saplings than on the surrounding shrubs. In synthesis, our work identifies the major biotic processes that appear to be favoring a community dominated by shrubs versus trees because seed dispersers, predators, and herbivores together favored shrub dispersal and establishment versus trees.     

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.     

Introduction of mammalian seed predators and the loss of an endemic flightless bird impair seed dispersal of the New Zealand tree Elaeocarpus dentatus

Understanding the mutualistic services provided by species is critical when considering both the consequences of their loss or the benefits of their reintroduction. Like many other Pacific islands, New Zealand seed dispersal networks have been changed by both significant losses of large frugivorous birds and the introduction of invasive mammals. These changes are particularly concerning when important dispersers remain unidentified. We tested the impact of frugivore declines and invasive seed predators on seed dispersal for an endemic tree, hinau Elaeocarpus dentatus, by comparing seed dispersal and predation rates on the mainland of New Zealand with offshore sanctuary islands with higher bird and lower mammal numbers. We used cameras and seed traps to measure predation and dispersal from the ground and canopy, respectively. We found that canopy fruit handling rates (an index of dispersal quantity) were poor even on island sanctuaries (only 14% of seeds captured below parent trees on islands had passed through a bird), which suggests that hinau may be adapted for ground‐based dispersal by flightless birds. Ground‐based dispersal of hinau was low on the New Zealand mainland compared to sanctuary islands (4% of seeds dispersed on the mainland vs. 76% dispersed on islands), due to low frugivore numbers. A flightless endemic rail (Gallirallus australis) conducted the majority of ground‐based fruit removal on islands. Despite being threatened, this rail is controversial in restoration projects because of its predatory impacts on native fauna. Our study demonstrates the importance of testing which species perform important mutualistic services, rather than simply relying on logical assumptions.     

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.