Joint evolution of mutualistic interactions, pollination, seed dispersal mutualism, and mycorrhizal symbiosis in trees.

Mycorrhizal symbiosis, seed dispersal, and pollination are recognized as the most prominent mutualistic interactions in terrestrial ecosystems. However, it remains unclear how these symbiotic relationships have interacted to contribute to current plant diversity. We analyzed evolutionary relationships among mycorrhizal type, seed dispersal mode, and pollination mode in two global databases of 699 (database I) and 10 475 (database II) tree species. Although database II had been estimated from phylogenetic patterns and therefore had lower certainty of the mycorrhizal type than database I, whose mycorrhizal type was determined by direct observation, database II allowed analysis of many more taxa from more regions than database I. We found evidence of joint evolution of all three features in both databases. This result is robust to the effects of both sampling bias and missing taxa. Most arbuscular mycorrhizal-associated trees had endozoochorous (biotic) seed dispersal and biotic pollination, with long dispersal distances, whereas most ectomycorrhizal-associated trees had anemochorous (abiotic) seed dispersal and wind (abiotic) pollination mode, with shorter dispersal distances. These results provide a novel scenario in mutualistic interactions, seed dispersal, pollination, and mycorrhizal symbiosis types, which have jointly evolved and shaped current tree diversity and forest ecosystem world-wide.

Seeds in the guts: can seed traits explain seed survival after being digested by wild ungulates?

Plants inhabiting open landscapes are often dispersed by ungulates and are expected to be adapted to this type of dispersal through their seed traits. To find which traits help seeds survive the passage through digestion of wild ungulates, we conducted a comprehensive feeding experiment with almost forty species of plants and three species of ungulates. We fed specified numbers of seeds to the animals, collected the dung, and germinated the dung content. We explored whether seed morphological traits and seed nutrient contents are good predictors of seed survival after passage through the ungulate digestive system. We also tested how the seed survival differed after the passage through different ungulate species. To find answers, we used GLMM with beta-binomial distribution and animal and plant species as random factor, respectively. We found that species survival and germination success were negatively correlated to seed elongation and the thickness of the seed coat. Even though phylogenetically correct GLMM did not yield significant results, when we tested species from commonly represented families, separately (legumes and grasses compared to all other species) different traits had statistically significant effects. In the case of seed elongation, the effect changed direction from negative to positive when legumes and grasses were left out. Our results suggest that seed traits enabling species survival after passage through the digestive tract are strongly phylogenetically conserved and different groups of plants evolved different ways of adapting to grazing pressure and utilize it for dispersal.

Seed dispersal function of the brown bear Ursus arctos on Hokkaido Island in northern Japan: gut passage time, dispersal distance, germination, and effects of remaining pulp.

Megafauna are important seed dispersers because they can disperse large quantities of seeds over long distances. In Hokkaido, Japan, the largest terrestrial animal is the brown bear (Ursus arctos) and other megafauna seed dispersers are lacking. Thus, brown bears are expected to have an important function as seed dispersers in Hokkaido. In this study, we, for the first time, evaluated the seed dispersal function of brown bears in Hokkaido using three fleshy-fruited trees and studied: (1) gut passage time (GPT) in feeding experiments, (2) seed dispersal distance using tracking data of wild bears, and (3) the effect of gut passage and pulp removal on germination rate. Most seeds were defecated intact, and less than 6% were broken. The average GPT without pulp was 3 h and 56 min to 6 h and 13 min, depending on the plant and trial. Each plant's average simulated seed dispersal distance was 202-512 m. The dispersal distance of Actinidia arguta seeds with pulp was significantly longer than those without pulp because of their longer GPT. The germination rate of defecated seeds without pulp was 19-51%, depending on the plant, and was significantly higher or not different comparing with that of seeds with pulp. We concluded that brown bears in Hokkaido are effective seed dispersers. In managing brown bears in Hokkaido, such ecological functions should be considered along with conserving the bear population and reducing human-bear conflicts.

Experimental evidence of dispersal of invasive cyprinid eggs inside migratory waterfowl.

Fish have somehow colonized isolated water bodies all over the world without human assistance. It has long been speculated that these colonization events are assisted by waterbirds, transporting fish eggs attached to their feet and feathers, yet empirical support for this is lacking. Recently, it was suggested that endozoochory (i.e., internal transport within the gut) might play a more important role, but only highly resistant diapause eggs of killifish have been found to survive passage through waterbird guts. Here, we performed a controlled feeding experiment, where developing eggs of two cosmopolitan, invasive cyprinids (common carp, Prussian carp) were fed to captive mallards. Live embryos of both species were retrieved from fresh feces and survived beyond hatching. Our study identifies an overlooked dispersal mechanism in fish, providing evidence for bird-mediated dispersal ability of soft-membraned eggs undergoing active development. Only 0.2% of ingested eggs survived gut passage, yet, given the abundance, diet, and movements of ducks in nature, our results have major implications for biodiversity conservation and invasion dynamics in freshwater ecosystems.

The important role of animal social status in vertebrate seed dispersal.

Seed dispersal directly affects plant establishment, gene flow and fitness. Understanding patterns in seed dispersal is, therefore, fundamental to understanding plant ecology and evolution, as well as addressing challenges of extinction and global change. Our ability to understand dispersal is limited because seeds may be dispersed by multiple agents, and the effectiveness of these agents can be highly variable both among and within species. We provide a novel framework that links seed dispersal to animal social status, a key component of behaviour. Because social status affects individual resource access and movement, it provides a critical link to two factors that determine seed dispersal: the quantity of seeds dispersed and the spatial patterns of dispersal. Social status may have unappreciated effects on post-dispersal seed survival and recruitment when social status affects individual habitat use. Hence, environmental changes, such as selective harvesting and urbanisation, that affect animal social structure may have unappreciated consequences for seed dispersal. This framework highlights these exciting new hypotheses linking environmental change, social structure and seed dispersal. By outlining experimental approaches to test these hypotheses, we hope to facilitate studies across a wide diversity of plant-animal networks, which may uncover emerging hotspots or significant declines in seed dispersal.

Fleshy fruit traits and seed dispersers: which traits define syndromes?

BACKGROUND AND AIMS: Fruit traits and their inter-relationships can affect foraging choices by frugivores, and hence the probability of mutualistic interactions. Certain combinations of fruit traits that determine the interaction with specific seed dispersers are known as dispersal syndromes. The dispersal syndrome hypothesis (DSH) states that seed dispersers influence the combination of fruit traits found in fruits. Therefore, fruit traits can predict the type of dispersers with which plant species interact. Here, we analysed whether relationships of fruit traits can be explained by the DSH. To do so, we estimated the inter-relationships between morphological, chemical and display groups of fruit traits. In addition, we tested the importance of each trait group defining seed dispersal syndromes. METHODS: Using phylogenetically corrected fruit trait data and fruit-seed disperser networks, we tested the relationships among morphological, chemical and display fruit traits with Pearson's correlations and phenotypic integration indices. Then, we used perMANOVA to test if the fruit traits involved in the analysis supported the functional types of seed dispersers. KEY RESULTS: Morphological traits showed strong intragroup relationships, in contrast to chemical and display traits whose intragroup trait relationships were weak or null. Accordingly, only the morphological group of traits supported three broad seed disperser functional types (birds, terrestrial mammals and bats), consistent with the DSH. CONCLUSIONS: Altogether, our results give some support to the DSH. Here, the three groups of traits interacted in different ways with seed disperser biology. Broad functional types of seed dispersers would adjust fruit consumption to anatomical limitations imposed by fruit morphology. Once this anatomical filter is sovercome, seed dispersers use almost all the range of variation in chemical and display fruit traits. This suggests that the effect of seed dispersers on fruit traits is modulated by hierarchical decisions. First, morphological constraints define which interactions can actually occur; subsequently, display and composition determine fruit preferences.

Statistical Comparison of Trait-Dependent Biogeographical Models Indicates That Podocarpaceae Dispersal Is Influenced by Both Seed Cone Traits and Geographical Distance.

The ability of lineages to disperse long distances over evolutionary timescales may be influenced by the gain or loss of traits adapted to enhance local, ecological dispersal. For example, some species in the southern conifer family Podocarpaceae have fleshy cones that encourage bird dispersal, but it is unknown how this trait has influenced the clade's historical biogeography, or its importance compared with other predictors of dispersal such as the geographic distance between regions. We answer these questions quantitatively by using a dated phylogeny of 197 species of southern conifers (Podocarpaceae and their sister family Araucariaceae) to statistically compare standard, trait-independent biogeography models with new BioGeoBEARS models where an evolving trait can influence dispersal probability, and trait history, biogeographical history, and model parameters are jointly inferred. We validate the method with simulation-inference experiments. Comparing all models, those that include trait-dependent dispersal accrue 87.5% of the corrected Akaike Information Criterion (AICc) model weight. Averaged across all models, lineages with nonfleshy cones had a dispersal probability multiplier of 0.49 compared with lineages with fleshy cones. Distance is included as a predictor of dispersal in all credible models (100% model weight). However, models with changing geography earned only 22.0% of the model weight, and models submerging New Caledonia/New Zealand earned only 0.01%. The importance of traits and distance suggests that long-distance dispersal over macroevolutionary timespans should not be thought of as a highly unpredictable chance event. Instead, long-distance dispersal can be modeled, allowing statistical model comparison to quantify support for different hypotheses.

The role of mucilage envelope in the endozoochory of selected plant taxa.

Myxodiaspory (formation of mucilage envelope around seeds and fruits) is a common adaptation to dry habitats known in many families of Angiosperms. The mucilage envelope of some seeds seems to be also a unique morphological adaptation which protects myxospermatic diaspores while passing through the bird's digestive system. To evaluate the protective potential of mucilage, we fed the diaspores of seven plant species (representing three different mucilage types and three species of non-mucilaginous plants) to pigeons, Columba livia domestica. Twenty-four hours later, we collected the droppings of pigeons and examined a total of 18,900 non-destroyed diaspores to check for mucilage presence and germination ability. Out of all the examined diaspores, 4.5% were mucilaginous seeds. Among them, the highest number (12.2-13.5%) of viable diaspores belonged to the hemicellulosic type of mucilage (from Plantago species). Only 3.7% of germinating diaspores with pectic mucilage (Linum usitatissimum) were collected, and no seeds representing cellulosic mucilage (e.g., Ocimum basilicum). For non-mucilaginous plants, we collected only a few individual seeds (0.1% out of 8100 seeds used). We noted that the mucilaginous seeds found in the droppings were able to germinate; however, the germination ability was generally smaller in comparison to the control (i.e., not digested) seeds. Our results revealed that the presence of mucilage envelope has an impact on diaspore dispersal and survivability. With our experiments, we demonstrated for the first time that the mucilage envelope, especially of the non-cellulosic type, supports endozoochory. We also showed that non-mucilaginous seeds can be occasionally dispersed via endozoochory and are able to germinate. The results of our studies can explain the ways of plants distribution at a small, local scale as well as in long-distance dispersal, e.g., between islands or even continents.

Experimental evaluation of the potential for crayfish plague transmission through the digestive system of warm-blooded predators.

The crayfish plague pathogen (Aphanomyces astaci) can be transmitted through the digestive system of fish, but its dispersal through mammalian and bird digestive tracts has been considered unlikely, and direct experimental evidence remains scarce. We present a small-scale transmission experiment with European otter and American mink fed with infected crayfish, and experiments testing survival of cultures of five A. astaci strains at temperatures corresponding to those inside mammal and bird bodies. The pathogen was neither isolated from predator excrements nor transmitted to susceptible crayfish exposed to excrements. In agar-based artificial media, it occasionally survived for 15 min at 40.5°C and for 45 min at 37.5°C, but not so when incubated at those temperatures for 45 min and 75 min, respectively. The five tested strains differed in resistance to high temperatures, two (of genotype groups E and D) being more susceptible than other three (of groups A, B and D). Their survival to some extent varied when exposed to the same temperature after several weeks or months, suggesting that some yet-unknown factors may influence A. astaci resistance to temperature stress. Overall, we support the notion that passage through the digestive tract of warm-blooded predators makes A. astaci transmission unlikely.

First evidence for the joint dispersal of mycorrhizal fungi and plant diaspores by birds.

Seed dispersal allows plants to colonise new sites and escape from pathogens and intraspecific competition, maintaining plant genetic diversity and regulating plant distribution. Conversely, most plant species form mutualistic associations with arbuscular mycorrhizal (AM) fungi in a symbiosis established immediately after seed germination. Because AM fungi are obligate symbionts, using the same dispersal vector as their host should be highly advantageous for their survival, but the co-dispersal of seeds and AM fungal spores has never been confirmed. We aim to clarify the potential role of European birds, essential dispersers for many plant species, as co-dispersers of seeds and AM fungal spores. In total, 63 bird droppings with intact seeds were placed in sterilised soil and maintained for 4 months in a protected environment to avoid contamination. Additionally, 173 bird droppings and 729 gauze swabs used to clean birds' feet were inspected for AM fungal spores. Although no spores were detected by direct observation of these samples, seven Rubus ulmifolius seedlings obtained from four independent droppings of Erithacus rubecula and Sylvia melanocephala were colonised by AM fungi. Our results show that birds can effectively co-disperse viable seeds and AM fungal spores, potentially over long distances, providing a pivotal mechanism to understand the cosmopolitan distribution of AM fungi.

Independent recruitment of a novel seed dispersal system by camel crickets in achlorophyllous plants.

The seeds of most heterotrophic plants, commonly referred to as dust seeds, are typically dispersed in the air like dust particles. Therefore, little attention has been paid to how seeds of heterotrophic plants are dispersed, owing to the notion that wind dispersal is the dominant strategy. However, inconspicuous but fleshy, indehiscent fruit can be observed in distantly related plants that have independently evolved full heterotrophy. Here I investigated the seed dispersal system in three unrelated fully heterotrophic plants with fleshy, indehiscent fruits (Yoania amagiensis, Monotropastrum humile and Phacellanthus tubiflorus) by direct observation, a differential exclusion experiment of fruit feeders and investigation on seed viability through the digestive tract. The present study revealed that camel crickets are the major seed disperser in three achlorophyllous plants in the study population. This represents the first evidence of seed dispersal by camel crickets in any angiosperm species. These heterotrophic plants grow in the understorey of densely vegetated forests where wind is probably an ineffective seed dispersal agent. Life-history traits of the achlorophyllous plants associated with heterotrophic lifestyles, such as colonization of dark understorey habitats and dust seeds, could facilitate independent recruitment of the novel endozoochorous seed dispersal system by camel crickets.

Effects of temporal variation in community-level fruit abundance on seed dispersal by birds across woody species.

PREMISE OF THE STUDY: In animal-dispersed plants, seed dispersal patterns of the same species in the same habitat can greatly differ among individuals and temporally. Many studies have revealed the pervasive effects of spatial variation in fruit abundance on seed dispersal of individual plants. By contrast, very few studies have investigated the temporal variation in seed dispersal. METHODS: We investigated the effects of conspecific and community-level fruit abundance on fruit removal rate and seed dispersal distance of six bird-dispersed woody species in a Japanese temperate forest for 3 yr. The six species share similar fruit characteristics but have different fruiting seasons. In addition, we also considered the effects of bird seasonality in diet and species composition on seed dispersal. KEY RESULTS: Inter-annual and inter-seasonal variation in community-level fruit abundance determined both fruit removal and the seed dispersal distance across species, but the effect differed with seasonal changes in bird behavior. Abundant fruit satiated fruit removal by birds only during fruit-feeding periods, not during insect-feeding periods. A scarcity of fruit increased the dispersal distance, but only during the migratory period. This difference was probably due to the different foraging behaviors of passing migrants and territorial birds. CONCLUSIONS: Our results illustrated that temporal variation in community-level fruit abundance could be a determining factor for seed dispersal in temperate forests. The effects of temporal variations in fruit abundance on seed dispersal patterns should be investigated in other ecosystems in order to understand their importance and associations with frugivore phenology.

Special delivery: scavengers direct seed dispersal towards ungulate carcasses.

Cadaver decomposition islands around animal carcasses can facilitate establishment of various plant life. Facultative scavengers have great potential for endozoochory, and often aggregate around carcasses. Hence, they may disperse plant seeds that they ingest across the landscape towards cadaver decomposition islands. Here, we demonstrate this novel mechanism along a gradient of wild tundra reindeer carcasses. First, we show that the spatial distribution of scavenger faeces (birds and foxes) was concentrated around carcasses. Second, faeces of the predominant scavengers (corvids) commonly contained viable seeds of crowberry, a keystone species of the alpine tundra with predominantly vegetative reproduction. We suggest that cadaver decomposition islands function as endpoints for directed endozoochory by scavengers. Such a mechanism could be especially beneficial for species that rely on small-scale disturbances in soil and vegetation, such as several Nordic berry-producing species with cryptic generative reproduction.

Whole angiosperms Wolffia columbiana disperse by gut passage through wildfowl in South America.

For the first time to our knowledge, we demonstrate that whole angiosperm individuals can survive gut passage through birds, and that this occurs in the field. Floating plants of the genus Wolffia are the smallest of all flowering plants. Fresh droppings of white-faced whistling duck Dendrocygna viduata ( n = 49) and coscoroba swan Coscoroba coscoroba ( n = 22) were collected from Brazilian wetlands. Intact Wolffia columbiana were recovered from 16% of D. viduata and 32% of Coscoroba samples (total = 164 plantlets). The viability of plants was tested, and asexual reproduction was confirmed. Wolffia columbiana is an expanding alien in Europe. Avian endozoochory of asexual angiosperm propagules may be an important, overlooked dispersal means for aquatic plants, and may contribute to the invasive character of alien species.

Strong among population variation in frugivory strength by functional diverse frugivores: a 'reciprocal translocation' experiment.

Fruit-frugivore interactions are critical for the dynamics and evolution of many plant communities. The strength of the interactions between a given plant species and different frugivore guilds (e.g., seed dispersers, seed predators) often vary in space due to changes in plant extrinsic factors (e.g., frugivore abundances) and plant traits (e.g., fruit size and reward). By reciprocally translocating Pyrus bourgaeana ripe fruits representative of five Mediterranean localities during 2 consecutive years, we experimentally quantified guild-specific variations among populations in frugivory strength, while accounted for plant-intrinsic and- extrinsic factors. Though overall fruit removal did not differ among localities, there were strong guild-specific differences in fruit removal strength. Fruit removal by pulp feeders, seed dispersers, and fruit predators varied among populations up to 8.5-, 5.6-, and 4.0-folds, respectively. These strong variations seemed mediated by changes in frugivore relative abundances rather than on availability of alternative fruits. As expected, all fruit traits considered (e.g., fruit size, pulp amount) markedly varied among tree populations. However, no frugivore guild showed preference for fruits from any locality, suggesting that fruit traits did not contribute much to differences in frugivory strength among populations. Since the functional diverse frugivore guilds played contrasting roles in P. bourgaeana dynamics (e.g., seed dispersal vs. seed predation), our study highlights the importance of accounting for functional diversity in frugivore guilds when estimating spatial variations in the strenght of seed dispersal. This investigation also illustrates a neglected but widely applicable experimental approach to identify the relative importance of extrinsic factors and fruit traits in mediating fruit-frugivore interactions.

The potential for dispersal of microalgal resting cysts by migratory birds.

Most microalgal species are geographically widespread, but little is known about how they are dispersed. One potential mechanism for long-distance dispersal is through birds, which may transport cells internally (endozoochory) and deposit them during, or in-between, their migratory stopovers. We hypothesize that dinoflagellates, in particular resting stages, can tolerate bird digestion; that bird temperature, acidity, and retention time negatively affect dinoflagellate viability; and that recovered cysts can germinate after passage through the birds' gut, contributing to species-specific dispersal of the dinoflagellates across scales. Tolerance of two dinoflagellate species (Peridiniopsis borgei, a warm-water species and Apocalathium malmogiense, a cold-water species) to Mallard gut passage was investigated using in vitro experiments simulating the gizzard and caeca conditions. The effect of in vitro digestion and retention time on cell integrity, cell viability, and germination capacity of the dinoflagellate species was examined targeting both their vegetative and resting stages. Resting stages (cysts) of both species were able to survive simulated bird gut passage, even if their survival rate and germination were negatively affected by exposure to acidic condition and bird internal temperature. Cysts of A. malmogiense were more sensitive than P. borgei to treatments and to the presence of digestive enzymes. Vegetative cells did not survive conditions of bird internal temperature and formed pellicle cysts when exposed to gizzard-like acid conditions. We show that dinoflagellate resting cysts serve as dispersal propagules through migratory birds. Assuming a retention time of viable cysts of 2-12 h to duck stomach conditions, cysts could be dispersed 150-800 km and beyond.

Great cormorants reveal overlooked secondary dispersal of plants and invertebrates by piscivorous waterbirds.

In wetland ecosystems, birds and fish are important dispersal vectors for plants and invertebrates, but the consequences of their interactions as vectors are unknown. Darwin suggested that piscivorous birds carry out secondary dispersal of seeds and invertebrates via predation on fish. We tested this hypothesis in the great cormorant (Phalacrocorax carbo L.). Cormorants regurgitate pellets daily, which we collected at seven European locations and examined for intact propagules. One-third of pellets contained at least one intact plant seed, with seeds from 16 families covering a broad range of freshwater, marine and terrestrial habitats. Of 21 plant species, only two have an endozoochory dispersal syndrome, compared with five for water and eight for unassisted dispersal syndromes. One-fifth of the pellets contained at least one intact propagule of aquatic invertebrates from seven taxa. Secondary dispersal by piscivorous birds may be vital to maintain connectivity in meta-populations and between river catchments, and in the movement of plants and invertebrates in response to climate change. Secondary dispersal pathways associated with complex food webs must be studied in detail if we are to understand species movements in a changing world.

Do deer and raccoons defecate in the right place? Fitness consequences of vertebrate seed dispersal for a deciduous forest herb.

Precision of seed placement in a heterogeneous environment is often assumed to select for the evolution of animal-mediated dispersal systems, but this hypothesis has rarely been tested in a multivariate sense. We quantify the microsite fitness benefits of dispersal by white-tailed deer (Odocoileus virginianus) and raccoons (Procyon lotor) for mayapple (Podophyllum peltatum), a shade-tolerant perennial herb, in deciduous forests of southeastern Ohio, USA. Micro-environmental variables were recorded at dung-deposition microsites, at rooting points of mayapple shoots, and at random (control) points in the forest. Fitness was assessed as the degree of overlap in ordinations of microsites by environmental variables. Mayapple occupied a broad sector (56%) of environment space corresponding to low and mid-slope positions, ravines, and proximity to trees. Deer and raccoon defecation placed dung in 71-74 and 86-95% of environment space, respectively, reaching mayapple microsites in 57-60 and 53-54% of cases. Deer placed dung in mayapple environment space 7-9% more often than predicted by random distribution, and raccoons placed dung in mayapple space 0-5% more often, consistent with only a modest degree of directed dispersal. Thus, the precision hypothesis is only weakly supported. The greatest fitness benefit of vertebrate dispersal appears to be the broad distribution of seeds, thereby increasing the probability of randomly reaching a suitable microsite. Imprecise dispersal suggests that secondary mechanisms of seed movement need to be explored in deciduous forest communities.

Overseas seed dispersal by migratory birds.

Long-distance dispersal (LDD) promotes the colonization of isolated and remote habitats, and thus it has been proposed as a mechanism for explaining the distributions of many species. Birds are key LDD vectors for many sessile organisms such as plants, yet LDD beyond local and regional scales has never been directly observed nor quantified. By sampling birds caught while in migratory flight by GPS-tracked wild falcons, we show that migratory birds transport seeds over hundreds of kilometres and mediate dispersal from mainland to oceanic islands. Up to 1.2% of birds that reached a small island of the Canary Archipelago (Alegranza) during their migration from Europe to Sub-Saharan Africa carried seeds in their guts. The billions of birds making seasonal migrations each year may then transport millions of seeds. None of the plant species transported by the birds occurs in Alegranza and most do not occur on nearby Canary Islands, providing a direct example of the importance of environmental filters in hampering successful colonization by immigrant species. The constant propagule pressure generated by these LDD events might, nevertheless, explain the colonization of some islands. Hence, migratory birds can mediate rapid range expansion or shifts of many plant taxa and determine their distribution.

Estimated Seed Shadow Generated by Japanese Martens(Martes melampus): Comparison with Forest-Dwelling Animals in Japan.

To evaluate the seed shadow generated by wild Japanese martens (Martes melampus), we combined data on their ranging behavior from the northern foot of Mt. Fuji, central Japan (seven males and three females) with data on gut passage time obtained from martens in Toyama Municipal Family Park Zoo (three males and one female). The movement distances varied, and mean distances for 0-1, 2-3, and 4-5 h intervals were 152.4, 734.7, and 1,162.4 m, respectively, with no significant sex difference. The mean gut passage time of ingested seeds was 7.4 h (range: 0.6-51.7 h), and two-thirds were defecated within 12 h. Seeds of fleshy fruits was frequently transported to 501-1,000 m, and 20% of ingested seeds were transported > 1,000 m from feeding sites. We found positive correlations between body size and home range of the animals in Japan and their seed dispersal distances. We conclude that Japanese martens are medium-range dispersers that can transport seeds from the source to open habitats conducive for germination and/or growth, partly due to scent marking behaviors.