The effects of kernel mass and nutrition reward on seed dispersal of three tree species by small rodents

In the present study, colored tags were fastened to individual seeds to investigate seed dispersal and seed fates of Pinus koraiensis, Corylus mandshurica, and Corylus heterophylla with different seed traits. Our aim was to determine the role of the proportion of kernel mass, caloric reward per seed, and tag color in affecting seed removal rates and seed fates. We predicted that higher proportion of kernel mass and caloric reward will favor seed removal and caching, while lower ones will facilitate seed consumption either in situ or after removal. Our results showed that the proportion of kernel mass, rather than seed size, played an important role in determining seed removal rates and seed fates. Seeds of C. heterophylla with the lowest proportion of kernel mass, regardless of their largest size and/or mass, had lower removal rates, lower level of caching but higher proportion of consumption in situ compared with P. koraiensis and C. mandshurica seeds. Seeds with higher caloric reward exhibited greater dispersal distances. Seed kernel proportion was closely correlated to seed removal rates and caching rates, while seed size and hull thickness seemed to play less important role in affecting seed dispersal, probably because the higher seed handling ability of small rodents in the experimental areas. Our investigation revealed no significant effect of tag color on seed removal rates and seed fates of the three seed species.     

Masting,seed dispersal and seed predation in the cycad Macrozamia communis

Summary Little is known about the adaptive value of mast seeding, a common phenomenon in temperate trees and shrubs. Masting is likely to affect both seed dispersal and seed predation. In systems where similar taxa of animals are involved in these two processes, the consequences of mast seeding are likely to be particularly complicated. This study examined the effects of mast seeding in a cycad, Macrozamia communis, on the dispersal of seeds, the pattern of dispersion of seeds and post-dispersal predation on seeds. Dispersal of seeds by possums was poorer from source plants in a masting population than from source plants in an adjacent, non-masting population. This resulted in fewer seeds per seeding female plant in the masting plot being dispersed to favourable sites. We conclude that this is caused by the feeding behaviour and movements of possums in the masting site. The abundance of seeds in this site did not satiate the post-dispersal predators, native rats. In fact, more seeds in this site were eaten than in the nonmasting site. We suggest that the mast seeding observed in M. communis may not be adaptive, but is more likely a consequence of other factors which synchromize flowering within local populations.     

The effects of seed abundance on seed predation and dispersal by rodents in <Emphasis Type="Italic">Castanopsis fargesii</Emphasis> (Fagaceae)

Based on the animal dispersal hypothesis and the predator satiation hypothesis, we examined the effects of seed abundance at both population (i.e., mast seeding) and community levels on seed predation and dispersal of Castanopsis fargesii (Fagaceae), a rodent-dispersed mast species in Eastern Asia. In a subtropical evergreen broadleaved forest in the Dujiangyan region of Sichuan Province, China, individual seeds with coded tin tags were tracked in two contrasting stands (seed-poor and seed-rich) over two years (2000, a low-seed year; 2001, a high-seed year). Our results showed that: (1) small rodents did not harvest the tagged seeds of C. fargesii more rapid in the high-seed year than in the low-seed year in either stand. But, seed harvest was significantly faster in the seed-rich stand than in the seed-poor stand. (2) The removal proportion was significantly lower in the high-seed year than in the low-seed year for either stand, but the removal proportion was slightly higher in the seed-poor stand than in the seed-poor stand. This indicates that high seed abundance decreases seed removal (predator satiation hypothesis). (3) There were only small differences about seed caching, seed survival and seedling establishment of C. fargesii between years and stands. During the survey, no cached seeds survived to geminate in the spring for both stands and years. (4) Mean dispersal distances of the cached seeds are much shorter in the high-seed year (3.1 m) than in the low-seed year (8.1 m) in the seed-rich stand, though similar trend is not examined in the seed-poor stand. Our results indicate that seed predation and dispersal of C. fargesii are influenced by both mast seeding and community-level seed abundance, which is not completely consistent with either the animal dispersal hypothesis or the predator satiation hypothesis, but seems more related to the predator satiation hypothesis.     

Effects of disperser abundance, seed type, and interspecific seed availability on dispersal distance

Seed dispersal distance is influenced by a variety of seed properties and functional responses of dispersers. However, to our knowledge, how and why seed dispersal distances are determined remains poorly understood. In the present study, seeds of sympatric tree species, Pinus koraiensis, Corylus mandshurica, Corylus heterophylla, and Quercus mongolica were released to investigate the effects of rodent abundance, seed type, and seed availability on seed dispersal. Our results showed that seeds of P. koraiensis were dispersed further than those of C. heterophylla and C. mandshurica regardless of the ambient rodent and seed abundances, reflecting a consistent effect of seed type on seed dispersal distances. Seed dispersal distance was greatly facilitated by lower per-capita seed abundance (the ratio of seeds to rodents); however, seed caching and cache survival were benefited from higher per-capita seed abundance. Although seed dispersal and seed caching of a particular tree species can be enhanced by its own seed availability, no consistent influence was detected at interspecific levels, reflecting different interspecific effects of seed availability on seed dispersal of sympatric seed species. Our results provide evidences that the effect of seed availability on seed dispersal should be evaluated in terms of per-capita seed abundance and interspecific effects, rather than the independent influence of seed or disperser abundances.     

Seed predation and dispersal of glabrous filbert (Corylus Heterophylla) and pilose filbert (Corylus Mandshurica) by small mammals in a temperate forest, northeast China

We investigated the seed dispersal of glabrous filbert (Corylus heterophylla) and pilose filbert (Corylus mandshurica), two large-seeded shrub species in a temperate forest, northeast China, September 2006. Small mammals such as Apodemus speciosus, Clethrlonomys rufocanus, and Eutamias sibiricus, were regarded as the main dispersal agents. More seeds were harvested by small mammals in pilose filbert (98%) than in glabrous filbert (87.5%) till our last survey. Seed removal rates differed between the two species. Fewer seeds of glabrous filbert (17.5%) were eaten in situ than pilose filbert (57.5%). More seeds of glabrous filbert were removed (70%), stay intact after removal (25.5%), eaten after removal (16%) than pilose filbert. However, more seeds were cached after removal in pilose filbert than in glabrous filbert (10.5 and 4%, respectively). Fewer tagged seeds of pilose filberts (14%) were missed than glabrous filberts (24.5%). About 8 and 12 primary caches were found in glabrous filbert and pilose filbert seeds respectively, indicating scatter hoarding. All of the removed seeds were distributed within 10 m of seed stations for both filberts. The average dispersal distances for glabrous filbert did not differ from pilose filbert. Only a small proportion of the caches remained till our last survey (2 and 1%, respectively). Based on the results, we found a difference in dispersal patterns of glabrous filbert and pilose filbert seeds. Evidences showed that glabrous filberts might be a less preferred seed species for small seed-eating mammals compared with pilose filbert, probably due to its harder and thicker husk and low seed profitability.     

Seed survival on experimental dishes in a central European old‐growth mixed‐species forest – effects of predator guilds,tree masting and small mammal population dynamics

Predation of tree seeds can be a major factor structuring plant communities. We present a three year study on tree seed survival on experimental dishes in an old‐growth forest in central Europe in Austria. We addressed species specific, spatial and temporal aspects of post‐dispersal seed predation. Seeds of Norway spruce Picea abies, European beech Fagus sylvatica, and silver fir Abies alba were exposed on dishes in different types of exclosures which allowed access only to specific guilds of seed predators. Removal experiments were carried out in two old‐growth forests and a managed forest (macro‐sites), including micro‐sites with and without cover of ground vegetation. We conducted the experiment in three consecutive years with a mast year of beech and spruce before the first year of the study. The seed removal experiments were combined with live trapping of small mammals being potential seed predators. Our experiments showed a distinctly different impact of different predator guilds on seed survival on the dishes with highest removal rates of seeds from dishes accessible for small mammals. We observed differing preferences of small mammals for the different tree species. Seed survival in different macro‐ and micro‐habitats were highly variable with lower seed survival in old growth forests. In contrast to our assumption, and in contrast to the satiation hypothesis which assumes higher seed survival in and directly after mast years, seed survival was lower in the year following the mast year of beech when a population peak of small mammals occurred and higher in intermast periods when subsequently small mammal population crashed. This suggests a higher importance of sporadic masting shortly after mast years in intermast periods for establishment of forest trees provided that pollination efficiency is high enough in such years. Combined with the high seed mortality observed after the mast year, this corroborates the important role of seed predation for forest dynamics. An altered synchrony or asynchrony of masting of different tree species and changed masting frequencies through climate change may thus lead to strong and non‐linear effects on forest dynamics.     

Seed dispersal of three sympatric oak species by forest rodents in the Qinling Mountains, Central China

Forest rodents play an essential role as seed dispersal vectors through their caching behaviors. Using seeds of Quercus aliena, Q. glandulifera, and Cyclobalanopsis engleriana (Fagaceae), which are dominant, but poorly studied species, in the Qinling Mountains, Central China, we investigated seed predation and dispersal by forest rodents in 2010 and 2011. There were significant differences in rodent seed-eating and caching strategies among the three tree species. Seeds of Q. aliena and C. engleriana had hard coats, high nutrition contents (e.g., protein, fat, and starch), and long germination schedules (C. engleriana only). They were less frequently eaten in situ, but more likely to be eaten after removal or cached. Seeds of Q. glandulifera had soft coats and low nutrition contents and were more often eaten in situ and less likely to be eaten after removal or cached. Our findings indicated that forest rodents were primarily responsible for seed predation and dispersal of these three tree species in the Qinling Mountains, and seed traits, especially coat hardness, nutrition content, and germination schedule, were important factors influencing rodent eating and caching behaviors. In addition, seed dispersal process of each tree species differed significantly between the 2 years, reflecting the effect of mast seeding on the eating and caching strategies of forest rodents.     

Effects of seed abundance on seed scatter-hoarding of Edward’s rat (<Emphasis Type="Italic">Leopoldamys edwardsi</Emphasis> Muridae) at the individual level

Mast seeding is a common phenomenon, and has important effects on seed dispersal and hoarding by animals. At population level, the predator satiation hypothesis proposes that the satiating effect of a large amount of seeds on a relatively small number of predators benefits seed survival in mast-seeding years. However, the effect of mast seeding on the scatter-hoarding of rodents at the individual level is largely unknown. In this study, we investigated the effects of seed abundance (by simulating mast seeding and non-mast seeding) on the removal, consumption and scatter-hoarding of seeds of Camellia oleifera (Theaceae) by Edward’s rat Leopoldamys edwardsi (Muridae) in seminatural enclosures in southwest China. We wanted to test the masting-enhanced hoarding hypothesis, which suggests that rodents tend to scatter-hoard more seeds in mast-seeding years in order to occupy more food resources. Our results indicate that L. edwardsi tended to disperse and scatter-hoard more seeds of C. oleifera per night with increasing seed abundance, and to eat less seeds per night when there was a high level of seed abundance in the enclosure experiments. These results support the masting-enhanced hoarding hypothesis. This capacity of rodents may be an evolutionary adaptation to the mast-seeding phenomenon. Our results suggest that mast seeding benefits forest regeneration not only through the predator satiation effect at the population level, but also through increased hoarding by animals at the individual level. Hongmao Zhang and Jinrui Cheng have contributed equally to this work.     

Endocarp thickness affects seed removal speed by small rodents in a warm-temperate broad-leafed deciduous forest, China

Seed traits are important factors affecting seed predation by rodents and thereby the success of recruitment. Seeds of many tree species have hard hulls. These are thought to confer mechanical protection, but the effect of endocarp thickness on seed predation by rodents has not been well investigated. Wild apricot (Prunus armeniaca), wild peach (Amygdalus davidiana), cultivated walnut (Juglans regia), wild walnut (Juglans mandshurica Maxim) and Liaodong oak (Quercus liaotungensis) are very common tree species in northwestern Beijing city, China. Their seeds vary greatly in size, endocarp thickness, caloric value and tannin content. This paper aims to study the effects of seed traits on seed removal speed of these five tree species by small rodents in a temperate deciduous forest, with emphasis on the effect of endocarp thickness. The results indicated that speed of removal of seeds released at stations in the field decreased significantly with increasing endocarp thickness. We found no significant correlations between seed removal speed and other seed traits such as seed size, caloric value and tannin content. In seed selection experiments in small cages, Père David's rock squirrel (Sciurotamias davidianus), a large-bodied, strong-jawed rodent, selected all of the five seed species, and the selection order among the five seed species was determined by endocarp thickness and the ratio of endocarp mass/seed mass. In contrast, the Korean field mouse (Apodemus peninsulae) and Chinese white-bellied rat (Niviventer confucianus), with relatively small bodies and weak jaws, preferred to select small seeds like acorns of Q. liaotungensis and seeds of P. armeniaca, indicating that rodent body size is also an important factor affecting food selection based on seed size. These results suggest endocarp thickness significantly reduces seed removal speed by rodents and then negatively affects dispersal fitness of seeds before seed removal of tree species in the study region. However, effect of endocarp thickness on final dispersal fitness needs further investigation because it may increase seed caching and survival after seed removal.     

Sensitivity to Seed Germination Schedule by Scatter‐Hoarding Pére David's Rock Squirrels During Mast and Non‐Mast Years

Mast seeding by oaks (Quercus) and other seed species has fundamental impacts on the behavior of individual scatter‐hoarding animals as well as their population and community dynamics. Scatter‐hoarding squirrels are highly sensitive to acorn germination schedule and stop germination by removing the embryo of non‐dormant acorns to prevent losses of energy and nutrients by rapid germination. However, we know little about how this behavior is affected by mast seeding. We investigated foraging decisions made by free‐ranging Pére David's rock squirrel (Sciurotamias davidianus) with three species that produce non‐dormant seeds and two species that produce dormant seeds (Fagaceae) during mast and non‐mast years in Central China from 2007–2010. Consistent with the food perishability hypothesis that squirrels make decisions to minimize the perishability of their caches, non‐dormant seeds were hoarded at a lower rate, but had a higher probability of embryo removal than dormant seeds. Compared with non‐mast years, a lower proportion of seeds were harvested during mast years, but a higher proportion of the harvested seeds were hoarded. In addition, the probability of embryo removal in the hoarded seeds was higher during mast years. Squirrels also dispersed seeds at shorter distances during mast years. Moreover, the interactions between mast seeding and seed germination schedule had significant effects on squirrel foraging decisions, including embryo removal. This study is the first to demonstrate that squirrels show greater sensitivity to seed germination potential when seeds are abundant.     

Predator satiation and extreme mast seeding in 11 species of Chionochloa (Poaceae)

Variation in annual flowering effort is described for 16 long datasets from 11 species of Chionochloa (Poaceae) in New Zealand. All populations exhibited extreme mast seeding. The most variable species was C. crassiuscula (coefficient of variation, CV=3.02) over 26 years at Takahe Valley, Fiordland, which is the highest published CV we know of worldwide. The other populations also had high CVs (lowest CV=1.42, mean CV=1.84) which were higher than for other well‐studied genera such as Picea, Pinus and Quercus. There were also frequent years of zero flowering (mean across all populations was 37.2% zero years; maximum 53% for C. rubra and C. crassiuscula over 19 years) whereas zero years are rare in other published masting datasets.Flowering was highly synchronous among species within a site (mean r=0.886), and also (though significantly less so) among sites. Among sites, synchrony was not significantly higher within‐species (mean r=0.711) than between‐species (r=0.690). Warm summer temperatures led to heavy flowering the following summer. Flowering synchrony increased with increasing synchrony in local deseasonalised summer temperatures, and decreased with increasing distance between sites.Mast seeding has been shown in Chionochloa to reduce losses to specialist flower or seed predators. Among‐species synchrony may be adaptive if species share a common seed predator. Developing seeds of at least 10 Chionochloa species are attacked by larvae of an undescribed cecidomyiid. In Takahe Valley, where masting is most pronounced, cecidomyiids attacked all six Chionochloa species in all four years studied. Mean annual losses were almost constant (10.0 to 13.4%) while flowering effort varied 100‐fold. The invariant losses are consistent with other evidence that the cecidomyiid may have extended diapause, which would make it harder to satiate by mast seeding. We hypothesise that one possible factor favouring such extremely high levels of mast seeding in Chionochloa is that its seed predator is very hard to satiate.     

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.     

Masting uncoupling: mast seeding does not follow all mast flowering episodes in a dioecious juniper tree

Evolutionary selective forces, like predator satiation and pollination efficiency, are acknowledged to be major causes of masting (the variable, periodic and synchronic production of seeds in a population). However, a number of recent studies indicate that resources might also play an important role on shaping masting patterns. Dioecious masting species offer a privileged framework to study the role of resources on masting variation, since male and female plants often experience different reproductive costs and selective pressures. We followed masting and reproductive investment (RI) of the dioecious tree Juniperus thurifera in two populations along 10 years and studied the different response of males and females to experimentally increased water and nutrient availability in a third population. Juniperus thurifera females invested in reproduction three times more resources than males. Such disparity generated different resource‐use strategies in male and female trees. Tree‐ring growth and water use efficiency (WUE) confirmed that sexes differed in their resource investment temporal pattern, with males using current resources for reproduction and females using resources accumulated during longer periods. Watered and fertilized female trees presented significantly higher flowering reproductive investments than males and experienced an extraordinary mast‐flowering event. However, seeding RI and mast seeding were not affected by the treatment. This suggests that although resource availability affects the reproductive output of this species, there are other major forces regulating masting on J. thurifera. During 10 years, J. thurifera male and female trees presented high and low flowering years more or less synchronously. However, not all mast flowering episodes resulted in mast seeding, leading to masting uncoupling between flowering and seeding. Since flowering costs represent only 1% of females’ total reproductive investments, masting uncoupling could be a beneficial bet‐hedging strategy to maximize reproductive output in spite of unpredictable catastrophic events.     

Effects of mast seeding and rodent abundance on seed predation and dispersal of <Emphasis Type="Italic">Quercus aliena</Emphasis> (Fagaceae) in Qinling Mountains,Central China

Rodents act as seed predators and dispersers and play an important role in the regeneration of plants. Seed production and rodent abundance may influence caching rodents’ decision to consume or cache seeds. We studied how did seed production and rodent abundance co-influence seed predation and dispersal by rodents in the mast seed years (2011 and 2013) and non-mast seed years (2012 and 2014) in the Qinling Mountains of Shaanxi Province, Central China. We found that: (1) The seed removal rates were much faster in the non-mast seed years than in the mast seed years. (2) Although the total number of seeds consumed (including eaten in situ and after removal) was higher in the non-mast seed years than in the mast seed years, no significant differences were found in the seeds that were eaten in situ and after removal among the 4 years. (3) Significant differences were observed in seeds that were cached among the 4 years, and more seeds were cached in the non-mast seed years than in the mast seed years. (4) The primary and secondary seed dispersal distances (including cached and eaten) were all longer in the non-mast seed years than in the mast seed years. Overall, these results indicate that non-mast seeding with low per capita seed availability could increase both seeds consumption and caching by rodents. Our results are partially supporting the predator satiation hypothesis.     

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.     

Interactions between ants and seeds of two myrmecochorous plant species in recently burnt and long‐unburnt forest sites

Myrmecochory (seed dispersal by ants) is a common seed dispersal strategy of plants in fire‐prone sclerophyll vegetation of Australia, yet there is little understanding of how fire history may influence this seed dispersal mutualism. We investigated the initial fate of seeds of two myrmecochorous plant species, the small‐seeded Pultenaea daphnoides J.C. Wendl. and the large‐seeded Acacia pycnantha Benth., in replicated burnt (3.25 years since fire) and unburnt (53 years since fire) forest plots in the Mount Lofty Ranges, South Australia. Specifically we measured (i) seed removal rates; (ii) the frequency of three ant–seed interactions (seed removal, elaiosome robbery and seed ignoring); (iii) the relative contribution of different ant species to ant–seed interactions; and (iv) the abundance of common interacting ant species. Rates of seed removal from depots and the proportion of seeds removed were higher in recently burnt vegetation and the magnitude of these effects was greater for the smaller‐seeded P. daphnoides. The overall proportion of elaiosomes robbed was higher in unburnt vegetation; however, the decrease in elaiosome robbery in burnt vegetation was greater for P. daphnoides than for A. pycnantha. Ants ignored seeds more frequently in burnt vegetation and at similar rates for both seed species. In total, 20 ant species were observed interacting with seeds; however, three common ant species accounted for 66.3% of ant–seed interactions. Monomorium sydneyense almost exclusively robbed elaiosomes, Rhytidoponera metallica typically removed seeds and Anonychomyrma nr. nitidiceps showed a mix of the three behaviours towards seeds. Differences in the proportions of seeds removed, elaiosomes robbed and seeds ignored appeared to be largely driven by an increase in abundance of A. nr. nitidiceps and a decrease in abundance of M. sydneyense in burnt vegetation. Understanding how these fire‐driven changes in the initial fate of myrmecochorous seeds affect plant fitness requires further investigation.     

Influence of between-year variation in the density of Rhus <Emphasis Type="Italic">trichocarpa</Emphasis> fruits on the removal of fruit by birds

The fruiting phenology and fruit removal patterns of Rhus trichocarpa Miq. (Anacardiaceae) were investigated in a warm-temperate secondary forest in Japan. Mature fruits of this species are dispersed by birds. Effects of fruit display size and canopy openness on fruit removal were investigated in years with different fruit densities (i.e., masting and non-masting years). Moreover, effects of increased canopy openness during winter on fruit removal were also investigated. Seasonal patterns of fruit removal were quite different between masting and non- masting years. In the non-mast year, fruits were removed by birds soon after maturation in the summer. In contrast, in the mast year, fruits were removed gradually by birds from summer to winter. Moreover, the rate of fruit removal was greater for trees with a larger display size in the non-mast year, whereas that was greater for trees with greater canopy openness in the mast year. Canopy openness increased in winter, and fruit removal in winter was enhanced in trees with a more open canopy only in the non-mast year. These observations strongly suggest that avian dispersers became satiated in the mast year, whereas fruit removal was enhanced in the non-mast year. In the mast year, although many fruits were not dispersed until winter, they were neither depredated nor rotten, and the long period of fruit removal by birds may have enhanced fruit dispersal. The large abundance of seedlings and saplings of this species in Japanese secondary forests suggests that this fruiting strategy is beneficial.     

The evolutionary ecology of mast seeding in trees

The hypothesis that masting by trees is a defensive strategy which satiates seed predators in mast years and starves them in the intervening periods is tested in 59 sets of data on the seed production and pre-dispersal seed-predation of 25 tree species. Twenty-lour of the 59 data-sets support the hypothesis and show a statistically significant positive relationship between the proportion of seeds surviving the pre-dispersal stage and the log₁₀ of the crop size for the same year. Evidence that pre-dispersal seed survival increases with the length of the mast interval is poor.
A positive relationship between the strength of the masting habit and the maxintum observed pre-dispersal seed mortality in a sample of 15 tree species suggests that the masting habit is best developed in predator-prone species. A survey of seed crop frequencies in the woody plant flora of Nortli America shows masting species to be under-represented amongst shrubs and amongst trees which disperse their seeds in fleshy dispersal units. The selection pressures and evolutionary constraints which operate on the evolution of masting plants and their seed predators are discussed.     

Post‐Dispersal Seed Removal and Germination Selected Tree Species Dispersed by Cebus capucinus on Barro Colorado Island,Panama1

Dispersal quality, an important component of seed disperser effectiveness, may strongly affect the rate of plant recruitment. Here we evaluated the quality of Cebus monkey dispersal by comparing the secondary removal fate and germination of fresh and Cebus‐ingested seeds of nine tree species on Barro Colorado Island, Panama. Overall, rates of secondary seed removal by vertebrates were low, with most Cebus defecations remaining undisturbed for extended periods on the forest floor. Only four of 30 feces were completely buried by dung beetles, and we found significantly higher vertebrate removal of defecated seeds than control seeds for only one species, Cordia bicolor. Seed germination varied greatly between plant taxa. Seeds of 3 out of 9 species showed significantly higher percent germination after monkey gut passage than control fresh seeds. Germination times tended to be shorter for defecated than for control seeds but were significantly different only for one of nine species, Cecropia insignis. Low rates of seed removal from Cebus feces, coupled with high germination probabilities, suggest high dispersal effectiveness for Cebus and contrasts strongly with patterns of post‐dispersal seed fate recorded for other primate species.     

Rodent handling of Araucaria araucana seeds

In an intermast year of very low seed production, we studied seed handling in an Araucaria araucana (Araucariaceae) forest in Neuquén Province, Argentina. Rodents identified in 844 photographs from automatic cameras removed 589 seeds marked with small embedded magnets. Within 12 days of removal, 460 were recovered using a magnetic field locator; 79% of recoveries were from burrows and caches in litter or soil. Rodents exhibited a diversity of seed‐handling strategies and differed in their potential as dispersal agents. Seed removal was most likely between 15.00 hours and 21.00 hours, but diurnal and nocturnal visits were recorded for all species. Oligoryzomys longicaudatus (Cricetidae) was a seed predator that took 8% of marked seeds and left none whole. Chelemys macronyx (Cricetidae) and Rattus norvegicus (Muridae) removed 25% and 19% of marked seeds respectively. These two species deposited the majority of seeds in groups of 10 or more in burrow larders that were unfavourable seedling establishment sites far from daylight. Abrothrix longipilis (Cricetidae) removed 43% of seeds, scatter‐hoarded the largest percentage of whole seeds (37%), moved some seeds farther than 40 m, and left them in favourable seedling establishment sites near daylight. For all species, the number and proportion of seeds cached whole increased as more seeds were removed. Rodents, especially A. longipilis, may play an important role in regeneration of A. araucana. Seed‐handling strategies and potentially effective dispersal are discussed in terms of masting seed production.