Abrupt increase in phosphorus and potassium fluxes during a masting event in a Bornean tropical forest

Mechanisms by which the productivity of tropical ecosystems is limited by nutrients is a long-standing question, but little information is available on the nutrient dynamics supporting the masting phenomenon in Southeast Asian evergreen rainforests. In this study we examined the nutrient sink and potential nutrient sources of masting in a Bornean tropical forest. We investigated if nutrient flux in fine litter, tree stems, and soils changed temporally in response to intense flower and fruit production. Fifty-five litter traps were installed in a 2-ha plot at the onset of flowering (April 2010), and litter and nutrient fluxes were monitored for more than 4 years (May 2010–December 2014). Wood cores of trunks and coarse roots of abundant species (Shorea spp.) and soil samples were collected in May 2010, September 2010, and September 2011 (coinciding with peak flowering, peak fruiting, and 1 year after fruiting, respectively). The P and K fluxes in the total litter were significantly greater in the mast year (2010) than non-mast years, whereas the Mg, N, and Ca fluxes did not vary in relation to masting. In line with the nutrient fluxes, P and K concentrations in coarse roots of flowering individuals of S. multiflora decreased in September 2011. The present results suggest that tropical trees require extraordinary amounts of P and K for masting, and may retranslocate stored nutrients to meet the elevated nutrient demands for masting.     

Contagious dispersal of seeds of synchronously fruiting species beneath invasive and native fleshy‐fruited trees

In subtropical Australia, many native and invasive plant species rely on a shared suite of frugivores, largely birds, for seed dispersal. Many native plants fruit during summer in this region, whereas most invasive plants fruit during winter, thus providing the opportunity for contagious dispersal of seeds beneath synchronously fruiting species. We sampled invasive and native seed rain beneath the canopy of a native summer‐fruiting tree Guioa semiglauca and an invasive winter‐fruiting tree Cinnamomum camphora, in three study sites over the course of a year. In July, during peak fruiting season for C. camphora and other invasive species, seed rain of invasive species was higher beneath C. camphora than G. semiglauca. This was partly due to the invasive tree Ligustrum lucidum, whose seed rain was three times higher beneath C. camphora than beneath the native tree. In February, seed rain of native species was more abundant beneath the canopy of G. semiglauca than beneath C. camphora, despite the fact that C. camphora was also fruiting at this time. This was probably due to the larger fruit crop produced by G. semiglauca at this time of year. Our study provides evidence that the presence of invasive bird‐dispersed plants may facilitate contagious seed dispersal of other invaders, and likewise native species may facilitate seed spread of other native plants.     

Disease resistance in Arabidopsisthaliana increases the competitive ability and the predicted probability of long‐term ecological success under disease pressure

Evidence is mounting that flowering by the mast-fruiting Dipterocarpaceae in Southeast Asia is triggered by ENSO events such that seeds are dispersed at the end of ENSO droughts. These droughts induce substantial defoliation and mortality of canopy trees, producing a favorable environment for seedling recruitment in the forest understory. Therefore, seedling release following droughts may have selected for synchronized, supra-annual fruiting in these rain forests. Currently, mast fruiting in Southeast Asia is generally regarded as an evolutionary response to seed predation by nomadic vertebrates. Separating the two causes for mast fruiting, seedling release and predator satiation, may be difficult if they are coupled in nature by ENSO droughts. Nevertheless, if the cue for masting is environmental, then the post-ENSO seedling environment should be considered a potential cause for masting, and if it operates in conjunction with predator satiation, then it may have provided the initial stimulus for supra-annual synchrony in fruiting.     

Strong “bottom‐up” influences on small mammal populations: State‐space model analyses from long‐term studies

“Bottom‐up” influences, that is, masting, plus population density and climate, commonly influence woodland rodent demography. However, “top‐down” influences (predation) also intervene. Here, we assess the impacts of masting, climate, and density on rodent populations placed in the context of what is known about “top‐down” influences. To explain between‐year variations in bank vole Myodes glareolus and wood mouse Apodemus sylvaticus population demography, we applied a state‐space model to 33 years of catch‐mark‐release live‐trapping, winter temperature, and precise mast‐collection data. Experimental mast additions aided interpretation. Rodent numbers in European ash Fraxinus excelsior woodland were estimated (May/June, November/December). December–March mean minimum daily temperature represented winter severity. Total marked adult mice/voles (and juveniles in May/June) provided density indices validated against a model‐generated population estimate; this allowed estimation of the structure of a time‐series model and the demographic impacts of the climatic/biological variables. During two winters of insignificant fruit‐fall, 6.79 g/m² sterilized ash seed (as fruit) was distributed over an equivalent woodland similarly live‐trapped. September–March fruit‐fall strongly increased bank vole spring reproductive rate and winter and summer population growth rates; colder winters weakly reduced winter population growth. September–March fruit‐fall and warmer winters marginally increased wood mouse spring reproductive rate and September–December fruit‐fall weakly elevated summer population growth. Density dependence significantly reduced both species' population growth. Fruit‐fall impacts on demography still appeared after a year. Experimental ash fruit addition confirmed its positive influence on bank vole winter population growth with probable moderation by colder temperatures. The models show the strong impact of masting as a “bottom‐up” influence on rodent demography, emphasizing independent masting and weather influences; delayed effects of masting; and the importance of density dependence and its interaction with masting. We conclude that these rodents show strong “bottom‐up” and density‐dependent influences on demography moderated by winter temperature. “Top‐down” influences appear weak and need further investigation.     

Nitrogen storage dynamics are affected by masting events in Fagus crenata

It is generally assumed that the production of a large crop of seeds depletes stores of resources and that these take more than 1 year to replenish; this is accepted, theoretically, as the proximate mechanism of mast seeding (resource budget model). However, direct evidence of resource depletion in masting trees is very rare. Here, we trace seasonal and inter-annual variations in nitrogen (N) concentration and estimate the N storage pool of individuals after full masting of Fagus crenata in two stands. In 2005, a full masting year, the amount of N in fruit litter represented half of the N present in mature leaves in an old stand (age 190–260 years), and was about equivalent to the amount of N in mature leaves in a younger stand (age 83–84 years). Due to this additional burden, both tissue N concentration and individual N storage decreased in 2006; this was followed by significant replenishment in 2007, although a substantial N store remained even after full masting. These results indicate that internal storage may be important and that N may be the limiting factor for fruiting. In the 4 years following full masting, the old stand experienced two moderate masting events separated by 2 years, whilst trees in the younger stand did not fruit. This different fruiting behavior may be related to different “costs of reproduction” in the full masting year 2005, thus providing more evidence that N may limit fruiting. Compared to the non-fruiting stand, individuals in the fruiting stand exhibited an additional increase in N concentrations in roots early in the 2007 growing season, suggesting additional N uptake from the soil to supply resource demand. The enhanced uptake may alleviate the N storage depletion observed in the full masting year. This study suggests that masting affects N cycle dynamics in mature Fagus crenata and N may be one factor limiting fruiting.     

Phenology of Tree Species in Bolivian Dry Forests

Phenological characteristics of 453 individuals representing 39 tree species were investigated in two dry forests of the Lomerío region, Department of Santa Cruz, Bolivia. The leaf, flower, and fruit production of canopy and sub–canopy forest tree species were recorded monthly over a two–year period. Most canopy species lost their leaves during the dry season, whereas nearly all sub–canopy species retained their leaves. Peak leaf fall for canopy trees coincided with the peak of the dry season in July and August. Flushing of new leaves was complete by November in the early rainy season. Flowering and fruiting were bimodal, with a major peak occurring at the end of the dry season (August–October) and a minor peak during the rainy season (January). Fruit development was sufficiently long in this forest that fruiting peaks actually tended to precede flowering peaks by one month. A scarcity of fruit was observed in May, corresponding to the end of the rainy season. With the exception of figs (Ficus), most species had fairly synchronous fruit production. Most canopy trees had small, wind dispersed seeds or fruits that matured during the latter part of the dry season, whereas many sub–canopy tree species produced larger animal– or gravity–dispersed fruits that matured during the peak of the rainy season. Most species produced fruit annually. Lomerio received less rainfall than other tropical dry forests in which phenological studies have been conducted, but rainfall can be plentiful during the dry season in association with the passage of Antarctic cold fronts. Still, phenological patterns in Bolivian dry forests appear to be similar to those of other Neotropical dry forests.     

Factors affecting seed rain beneath fleshy-fruited plants

We investigated factors affecting seed rain beneath nine fleshy-fruited fruiting plant species growing in a 1-ha plot of planted Pinus thunbergii in central Japan. We tested whether the numbers of seeds and seed species dropped by birds beneath fruiting plants were correlated with the number of fruits removed by birds from the plants. Most of fruiting plant species with high fruit removal had significantly high seed rain. Both the numbers of seeds and seed species dropped were significantly, positively correlated with the number of fruits removed across for all fruiting plant species. Therefore, fruit removal predicted the difference among heterospecific fruiting plants in seed rain. We also tested whether the number of fruits removed from fruiting plants by birds was related with fruit crop size, fruit size, and height of the plants, and the numbers of fruits and fruit species of neighboring plants near the plants. Most of fruiting plant species with high fruit crop size had significantly high fruit removal. The number of fruits removed was significantly, positively correlated with both the fruit crop size and the number of neighboring fruits across the nine fruiting plant species. However, the effect of the neighboring fruit density on fruit removal was lower remarkably than that of fruit crop size. Therefore, fruit crop size best predicted the differences among heterospecific fruiting plants in fruit removal. We suggest that fruiting plant species with high fruit crop size and high fruit removal contribute to intensive seed rain beneath them. This revised version was published online in June 2006 with corrections to the Cover Date.     

Masting in a temperate tree: Evidence for environmental prediction?

Many plant species produce large fruit crops in some years and then produce few or no fruits in others. Synchronous, inter‐annual variation in plant reproduction is known as ‘masting’ and its adaptive significance has yet to be fully resolved. For 8 consecutive years, I quantified every fruit produced by 22 females of a New Zealand tree species (Dysoxylum spectabile), which has an unusual habit of taking a full calendar year to mature fruits after flowering. Fruit production varied strongly among years and was tightly synchronized among trees. Annual variability in fruit production declined with total reproductive output, indicating trees with lower fecundity exhibited a stronger tendency to mast. Although unrelated to temperature, annual fruit production was positively related to precipitation during annual periods of fruit development, and negatively related to fruit production in the previous year. Seedlings had higher rates of survivorship in a wet, high‐seed year than in a dry, low‐seed year, suggesting that seedlings might be drought sensitive. Therefore, D. spectabile produced large fruit crops during periods of high rainfall prior to fruit maturation, which may enhance survivorship of drought‐intolerant seeds. Results were inconsistent with several hypotheses that are widely believed to be the most likely explanations for masting. Instead, results were consistent with the environmental prediction hypothesis, suggesting that this hypothesis may be more important than previously appreciated.     

An increase in canopy cover leads to masting in Quercus ilex

Masting is the intermittent and synchronous production of large crops, but its relation to tree growth remains elusive despite the ecological relevance of mast seeding. The production of huge fruit crops has been linked to the accumulation and consumption of resources as nutrients and carbohydrates, but no conclusive assessment has supported this assumption. To evaluate if masting takes place once trees’ canopies reach maximum foliage, changes in canopy cover were measured in Quercus ilex susbp. ballota stands before and after a masting event using the normalized difference vegetation index (NDVI). The results on the whole underline that masting in Q. ilex occurred once maximum levels of NDVI and canopy cover were reached. After the masting event, NDVI dropped, leaf shedding increased and trees produced shorter shoots, narrower tree rings and fewer acorns than before the masting event. These findings support our contention that an increase in canopy cover precedes masting.     

Impact on fruit removal and seed predation of a secondary metabolite, emodin, in Rhamnus alaternus fruit pulp

There are two contradictory approaches to explaining the presence of secondary metabolites in ripe fruits. One holds that they evolved toward enhancing dispersal success (adaptive approach); the other claims that they evolved primarily to deter herbivores from eating leaves and seeds and that their presence in ripe fruits is a byproduct of that function (non‐adaptive approach). We tested the validity of three hypotheses of the adaptive approach that explain the presence of secondary metabolites in ripe fruits. We explored the current function of a secondary metabolite, emodin, in Mediterranean buckthorn (Rhamnus alaternus, Rhamnaceae) fruits by relating intraspecific variation and seasonal patterns of concentration to fruit removal and seed damage and by conducting feeding trials with captive birds presented artificial fruits that varied in emodin concentration. The concentration of emodin in the pulp of 10–13 Rhamnus plants from the same population was determined by HPLC every month during two fruiting seasons. Fruit removal by birds and seed predation by invertebrates and microbes were determined for the same plants. Emodin concentration rose during the first stages of ripening, reaching a peak before the fruits were ripe, and then decreased to a minimum when the fruits were ripe. No significant correlation between emodin concentration and ripe fruit removal rate among trees was observed in the first year, whereas in the second year the correlation was positive and significant. Thus, the impact of emodin on fruit selection varied between years, suggesting that emodin concentration does not solely govern fruit selection. A significant negative correlation was found in the first year between emodin concentration and seed predation during the first fruiting month. The yellow‐vented bulbul (Pycnonotus xanthopygos), a seed dispersing bird, distinguished between artificial foods that differed in emodin concentration (control, 0.001% and 0.002%), always preferring the lower concentration. In contrast, house sparrows, (Passer domesticus), a seed predator, did not detect such differences in emodin concentration but did distinguish between control foods and food with 0.005% and 0.001% emodin. We suggest that emodin has an ecological role, preventing seed predation by invertebrates and microbes without decreasing fruit removal by avian dispersers.     

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

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

Non‐continuous reproductive phenology of animal‐dispersed species in young forest restoration plantings

Tree species that produce resources for fauna are recommended for forest restoration plantings to attract pollinators and seed dispersers; however, information regarding the flowering and fruiting of these species during early growth stages is scarce. We evaluated the reproductive phenology of animal‐dispersed tree species widely used in Atlantic Forest restoration. We marked 16 animal‐dispersed tree species in 3‐ to 8‐year‐old forest restoration plantings in Itu‐São Paulo, southeast Brazil. We noted the age of the first reproductive event, flowering and fruiting seasonality, percentage of trees that reached reproductive stages, and intensity of bud, flower, and fruit production for each species. Flowering and fruiting are seasonal for most species; only two, Cecropia pachystachya and Ficus guaranitica, exhibited continuous flowering and fruiting throughout the year; we also identified Schinus terebinthifolia and Dendropanax cuneatus fruiting in the dry season during resource scarcity. Therefore, we recommend all as framework species, that is, species that are animal‐dispersed with early flowering and fruiting potential, for forest restoration. Further, we recommend identifying and planting similar animal‐dispersed tree species that produce fruits constantly or in the dry season to maximize fauna resource availability throughout the year in tropical forest restoration plantings. Abstract in Portuguese is available with online material     

Temporal variations in seed dispersal patterns of a bird-dispersed tree, <Emphasis Type="Italic">Swida controversa</Emphasis> (Cornaceae), in a temperate forest

The seed dispersal patterns of bird-dispersed trees often show substantial seasonal and annual variation due to temporal changes in frugivorous bird and bird-dispersed fruit distributions. Elucidating such variation and how it affects plant regeneration is important for understanding the evolution and seed dispersal maintenance strategies of these plants. In this study, we investigated the seed dispersal quantity and distance of a bird-dispersed plant, Swida controversa, for 2 years and detected large seasonal variations in dispersal pattern. Early in the fruiting season, short seed dispersal distance and large amounts of fruit consumption by birds (seed dispersal quantity) were observed. In contrast, late in the fruiting season, a long seed dispersal distance and small seed dispersal quantity were observed. This relationship between seed dispersal distance and quantity may help to maintain constant seed dispersal effectiveness during the long S. controversa fruiting season. Annual variation was also detected for both seed dispersal quantity and distance. More effective seed dispersal was achieved in the masting year, because both seed dispersal quantity and distance were greater than that in the non-masting year. These seed dispersal dynamics may contribute to the evolution and maintenance of S. controversa masting behavior. Thus, we identified substantial temporal variation on both seasonal and annual scales in the seed dispersal pattern of a bird-dispersed plant. The temporal variation in seed dispersal pattern revealed in this study probably plays a substantial role in the life history and population dynamics of S. controversa.     

Pollen coupling of forest trees: forming synchronized and periodic reproduction out of chaos

Many of the tree species in mature forests show masting; their reproductive activity has a large variance between years and is often synchronized between different individuals. In this paper, we analyse a globally coupled map model in which trees accumulate photosynthate every year, produce flowers when the energy reserve level exceeds a threshold, and set seeds and fruits at a rate limited by pollen availability. Without pollen limitation, the trees in the forest show independent chaotic fluctuation. Coupling of trees via pollen exchange results in reproduction being synchronized partially or completely over the forest. The whole forest shows diverse dynamical behaviors determined by the values of two essential parameters; the depletion coefficient k and the coupling strength beta. We find perfectly synchronized periodic reproduction, synchronized reproduction with a chaotic time series, clustering phenomena, and chaotic reproduction of trees without synchronization over individuals. There are many parameter windows in which synchronized reproduction of trees shows a stable periodic fluctuation. For perfectly synchronized forests, we can calculate all the Lyapunov exponents analytically. They show that synchronized reproduction of all the trees in the forest can only occur when trees flower at low (but positive) levels in a significant fraction of years, resulting in small fruit sets due to outcrossed pollen limitation. This is consistent with the observation that the distinction between mast years and non-mast years is often not clear cut.     

Fruiting phenology of animal-dispersed plants in response to winter migration of frugivores in a warm temperate forest on Yakushima Island, Japan

The fruiting phenology of animal-dispersed plants was observed in a warm temperate, evergreen forest on Yakushima Island. The number of ripe fruits was counted for 22 trees, four lianas and one parasitic epiphyte species with sapfruit. These fruits were consumed by birds and Japanese macaques (Macaca fuscata yakui). Birds with small gapes (e.g. Japanese white-eye [Zosterops japonica]) consumed only small fruit less than 6 mm in diameter, while birds with large gapes (e.g. red-capped green pigeon [Sphenurus formosae]) and Japanese macaques consumed a wide range of fruits from 4 to 16 mm in diameter. The larger animals did not ignore the smaller fruits. Brown-eared bulbul (Hypsipetes amaurotis) and Japanese white-eye were the main consumers of sapfruit in terms of frequency in winter. Some of the observed consumers were year-round residents, but most of the consumers migrated to Yakushima Island from the main islands of Japan to overwinter (from November to March), and their abundance in winter was four times as high as during the rest of the year (from May to October). In 23 of the 27 plant species investigated, sapfruit production coincided with their immigration season, whereas tree species bear capsules and nuts during autumn from September to November. We suggest that sapfruit species set their ripe period to the season when frugivorous birds are most abundant.     

Fruit characteristics and factors affecting fruit removal in a Panamanian community of strangler figs

We describe fruiting characteristics for 12 species in a community of strangler figs (Moraceae: Urostigma) studied in Panama. We quantify diurnal and nocturnal removal rates and proportions of fruits removed, and relate them to the activities of the main dispersers of the figs: bats and birds. These results combined with previous studies show that there are clear differences between fig species with fruit that ripen red and those with fruit that remain green(ish). In the red-fruited species, the fruit are small, ripen asynchronously over relatively long periods, produce little scent, and are mainly taken during the day by birds. In contrast, in the green(ish)-fruited species, the fruits are larger, span a range of sizes, ripen relatively synchronously, produce very distinctive aromas, and are mainly taken at night by bats. This dichotomy in fruiting characteristics suggests coadaptive links between groups of dispersers and different species within the genus Ficus. All fig species produce a range of fruit crop sizes (10–155 fuits/m² canopy area) of which a high proportion were removed by seed dispersers (>80%). Removal rates (fruit removed per day) were positively correlated with crop size, suggesting that trees with large crop size attract more frugivores. Removal rates of green-fruited figs were significantly lower and persistence and abortion of ripe fruit were significant higher around full moon, apparently due to the reduced activity of bats. We further estimate the number of bats that are sustained by a tree fruit crop and account for the observed fruit removal. We then discuss the evidence for coadaptation between different groups of figs and their seed dispersers, Finally, we consider the conservation implications for figs as keystone resources in tropical forests. Received: 26 April 1999 / Accepted: 10 January 2000     

An appraisal of seed enumeration and videographic techniques for determining seed removal rates by birds

We examined the efficacy of seed enumeration and videographic techniques for determining seed removal by birds from indigenous (Chrysanthemoides monilifera and Olea europaea subsp africana) and alien (Lantana camara and Solanum mauritianum) shrubs at different study sites in the Cape Floristic Region. The seed enumeration technique involved counting the numbers of fruits and associated seeds removed monthly by birds, excluding those naturally abscised, from the shrub canopy. The videographic technique involved visual counts from images of the numbers of fruits and associated seeds consumed by birds over specific time intervals captured by a digital camcorder. Daily seed removal rates by all birds, irrespective of species, measured by both techniques were similar with no significant interactions evident between measuring techniques, site and shrub species. Both techniques displayed higher seed removal from tiny‐seeded S. mauritianum than other shrub species; this was also evident among individual bird species. However, the seed enumeration technique was unable to discriminate between foraging organisms, contamination of traps by wind‐blown fruits abscised from neighbouring branches and fruit theft from the canopy and the traps. In contrast, the videographic technique provided permanent visual and time‐lapse records for individual foraging bird species allowing greater measurement precision and interpretation of fruit removal behaviour by birds. We recommend use of the videographic technique over the seed enumeration technique for studying vertebrates’ seed removal in a detailed manner.     

Fruit production in Sorbus aucuparia L. (Rosaceae) and pre-dispersal seed predation by the apple fruit moth (Argyresthia conjugella Zell.)

Variation in fruit production and pre-dispersal seed predation by Argyresthia conjugella was studied in␣four populations of Sorbus aucuparia in northern Sweden.␣The number of infructescences, fruits per infructescence, consumed seeds and developed unattacked seeds per fruit were scored in marked trees from 1984 to 1990. The results showed that the number of fruits produced in each population determined the number of seed predators occurring in the host population, as the yearly number of seed predators was significantly and positively correlated with yearly number of fruits, in all but one population. The seed predators showed a delay in response to variation in number of fruits produced. This lag in response resulted in a large proportion of fruits being attacked and seeds consumed in a bad fruiting year that followed a good fruiting year, and vice versa. The proportion of fruits attacked and seeds consumed was largest in the population showing the greatest between-year variation in fruit production and lowest in the population showing the lowest between-year variation in fruit production. Furthermore, the individuals within the former population were synchronised, while they were not in the latter population. These results contradict one of the possible explanations of mast-seeding, where large synchronised between-year variation is supposed to reduce pre-dispersal seed predation. Instead, differences in attraction of the seed predator to differences in fruit crop size could explain the observed difference in seed predation between the two populations with opposite fruiting patterns. Within each population, irrespective of year, the proportion of fruits attacked and seeds consumed was independent of a tree's fruiting display. Therefore, trees with high fruit production, despite harbouring the largest number of seed predators, produced the largest number of developed seeds in absolute numbers, compared to trees that produced few fruits. Received: 25 February 1996 / Accepted: 30 November 1996     

Seasonal variation in fleshy fruit use and seed dispersal by the Japanese black bear (<Emphasis Type="Italic">Ursus thibetanus japonicus</Emphasis>)

Fruit use by the Japanese black bear (Ursus thibetanus japonicus) and seed clumping in bear scat were studied in central Japan using fecal analyses. Between May and November 2003 and 2004, the life form and fruit size of plants consumed by bears and the species composition and intactness of seeds contained in scat were examined in five transects (approximately 10 km × 10 m) in broad-leaved deciduous forests. In 2003, scats with seeds were found only in the autumn, when fruiting trees and shrubs were abundant. In 2004, scats with seeds occurred intermittently from the summer, when fruiting plants were rare, up to the autumn. Yearly and seasonal variation in fruit use reflects the opportunistic foraging behavior of Japanese black bears. Seven of the nine plant species detected in scats had medium-sized fruits (6–15 mm width), whereas the other two species had relatively large fruits (20–100 mm width). In total, 14,492 seeds were detected, of which 97.6% were intact; the remainder were damaged. Intact seeds of one or two species were found in each scat. The number of intact seeds per scat ranged from 1 to 5476. Japanese black bears seldom digest ingested seeds, thereby contributing to the seed dispersal of their food plants, including species with fruits that are too large to be swallowed by frugivorous birds.     

Testing Broadcast Seeding Methods to Restore Urban Forests in the Presence of Seed Predators

Forest restoration in urban areas often occurs in isolation from remnant forest, limiting the chances for recolonization by native species. Plants with bird‐dispersed seeds can be particularly vulnerable to dispersal limitation and regeneration can be further impeded by non‐native seed predators. We used a factorial experiment to investigate broadcast seeding as a method to reintroduce trees with large seeds and fleshy fruits into early successional forests. We assessed rates of seed and fruit loss, germination and seedling establishment in three seed treatments: (1) caging to exclude introduced mammalian seed predators; (2) removal of fleshy fruit pericarp; and (3) placing seeds in nutritionally enriched clay balls. Across all species (Beilschmiedia tawa, Elaeocarpus dentatus, and Litsea calicaris) seeds and fruits accessible to mammalian predators suffered significantly greater loss (58%) than those protected by cages (4%). However, seed and fruit loss in the presence of predators was reduced to only 35% across all species by the treatment combining the removal of fruit flesh and clay ball application to seeds. Establishment of B. tawa seedlings after 1 year was significantly enhanced by the clay ball treatment (12% of seeds sown vs. 6% without clay balls). Very low establishment rates were recorded for E. dentatus and L. calicaris. Broadcast seeding was found to be a viable method of improving regeneration of large‐seeded late successional trees and may be a cost‐effective alternative to planting saplings. Seedling establishment can be improved with fruit flesh removal and clay ball treatments, especially in the presence of mammalian seed predators.