Seed Dispersal of a High Quality Fruit by Specialized Frugivores: High Quality Dispersal?1

Dispersal quality, as estimated by the cumulative effects of dispersal, germination, seed predation, and seedling survival, was examined for Beilschmiedia pendula (Lauraceae) in Monteverde, Costa Rica. I determined the pattern of dispersal by finding seeds deposited by birds, protected the seeds from seed predators with cages to assess germination and seedling survival, and examined seed predation rates with marked seeds. Seed predation, germination, and seedling survival were compared between seeds naturally dispersed by birds and seeds placed at randomly located sites. Approximately 70 percent of seeds dispersed by birds (N= 244) were deposited <10 m from crown edges of fruiting B. pendula trees, although some seeds were dispersed at least 70 m away. Larger seeds were more likely to be dispersed under or close to the parent trees, and larger seeds produced larger seedlings. Seed size was not correlated directly with seedling survival, but larger seedlings at three months were most likely to survive one year. Seed predation by mammals and insects and seedling mortality due to fungal pathogens were concentrated beneath the crowns of parent trees. Seedlings and saplings were more abundant beneath fruiting B. pendula trees, but individuals farther away were taller on average. Thus, dispersal is beneficial for B. pendula, but such benefits appear most pronounced at a small spatial scale; seeds dispersed >30 m from the crown edges actually had a lower probability of survival than those dispersed 10–20 m. Only 10 percent of B. pendula. seeds received high‐quality dispersal in terms of landing in the zone with the highest per seed probability of seedling survival 10–20 m from parental crowns.     

Linking seed dispersal,germination and seedling recruitment in the invasive species <Emphasis Type="Italic">Berberis darwinii</Emphasis> (Darwin’s barberry)

Seedling recruitment is a multi-phased process involving seed production, dispersal, germination, seedling establishment and subsequent survival. Understanding the factors that determine success at each stage of this process is of particular interest to scientists and managers seeking to understand how invasive species spread and persist, and identify critical stages for management. To understand the factors and processes influencing recruitment of the invasive species Berberis darwinii Hook. (Darwin’s barberry), temporal and spatial patterns of seed dispersal, germination and seedling establishment were examined. Seed dispersal from a large source population was measured over two fruiting seasons, and subsequent patterns of seedling emergence and survival within each cohort were measured. Seed longevity was tested under both natural and artificial conditions. Seeds were widely dispersed by birds, up to 450 m from the source population. Dispersal was essential to seedling establishment, as few seedlings survived beneath the parent canopy. Seeds were relatively short-lived in the soil under both field and glasshouse conditions, with few surviving for more than 1 year. Patterns of newly emerged seedlings largely reflected patterns of seed rain, but seedling survival was significantly affected by distance from source population, seedling density and light environment. These results suggest that recruitment of B. darwinii is dependent on dispersal of seeds to favourable microsites. Management priorities should include the removal of fruiting plants, and seedling control in highlight areas.     

Recruitment of two Opuntia species invading abandoned olive groves

In Europe, many agricultural areas are now abandoned and hence can be invaded by exotic species. The abundance and spatial distribution patterns of two Opuntia species were studied in old olive groves in the Parc Natural del Cap de Creus, Catalonia (Spain). Seedling recruitment (97.3% and 51.5% of juveniles for O. maxima and O. stricta, respectively) was higher than recruitment by cladodes. O. maxima had more seedlings recruited beneath olive trees and beneath Opuntia adults than expected. Most O. stricta seedlings were also located beneath Opuntia adult plants. However, although most seedlings were recruited beneath Opuntia, some (10–30%) were found away from putative parental plants. This may be due to seed dispersal by birds and wild boars. Seeds dispersed by wild boars were not significantly more viable than seeds from intact fruits. Seedlings grow very slowly but have a high survival rate. In conclusion, Opuntia seedling recruitment is very successful and ensures the persistence of these species within old olive groves. Consequently, it prevents restoration from an agricultural land-use back to the native community.     

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

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

Small Tent-Roosting Bats Promote Dispersal of Large-Seeded Plants in a Neotropical Forest

In Neotropical regions, fruit bats are among the most important components of the remaining fauna in disturbed landscapes. These relatively small-bodied bats are well-known dispersal agents for many small-seeded plant species, but are assumed to play a negligible role in the dispersal of large-seeded plants. We investigated the importance of the small tent-roosting bat Artibeus watsoni for dispersal of large seeds in the Sarapiquí Basin, Costa Rica. We registered at least 43 seed species > 8 mm beneath bat roosts, but a species accumulation curve suggests that this number would increase with further sampling. Samples collected beneath bat feeding roosts had, on average, 10 times more seeds and species than samples collected 5 m away from bat feeding roosts. This difference was generally smaller in small, disturbed forest patches. Species-specific abundance of seeds found beneath bat roosts was positively correlated with abundance of seedlings, suggesting that bat dispersal may influence seedling recruitment. Our study demonstrates a greater role of small frugivorous bats as dispersers of large seeds than previously thought, particularly in regions where populations of large-bodied seed dispersers have been reduced or extirpated by hunting.     

Cache site selection by chipmunks (Tamias spp.) and its influence on the effectiveness of seed dispersal in Jeffrey pine (Pinus jeffreyi)

The effectiveness of Jeffrey pine (Pinus jeffreyi) seed dispersal performed by seed-caching yellow pine chipmunks (Tamias amoenus) and lodgepole chipmunks (Tamias speciosus) was compared to that of wind dispersal in the Sierra Nevada of western Nevada. Wind-dispersed seeds typically fall under or near the parent tree. Chipmunks removed 90 and 97% of 1064 radioactive seeds from each of two simulated wind-dispersed seed shadows in less than 24 h. Wind-dispersed seeds were deployed within 12 m of the two source trees, but chipmunk caches were found from 2–69 m from the trees. Chipmunks carried nearly all seeds away from source trees, greatly reducing the density of seeds under and near source trees. Caches contained from 1–35 seeds and most were buried 7–21 mm deep. Chipmunks cached in open bitterbrush shrubland with mineral soils much more than expected and cached in closed-canopy Jeffrey pine and lodgepole pine forests with thick needle litter much less than expected. Many Jeffrey pine seedlings and saplings grow in the bitterbrush habitat and few grow in the pine forests. Ten and 20% of the original caches survived until April, the time of seed germination, at the two sites. The movement of wind-dispersed seeds is random relative to environmental variables important in seedling survival, and the wind in coniferous forests cannot quickly bury seeds. The quality of seed dispersal rendered by chipmunks was superior to that provided by the wind because the chipmunks quickly harvested seeds on the ground, moved them away from source trees, and buried them in the ground in habitats and microhabitats where they were more likely to establish new seedlings. The increased quality of seed dispersal provided by animals relative to the wind may help explain why over twenty species of pines have evolved seeds and cones that are adapted for dispersal by seed-caching animals.     

Is farther seed dispersal better? Spatial patterns of offspring mortality in three rainforest tree species with different dispersal abilities

The paradigm that tropical trees with farther seed dispersal experience lower offspring mortality is currently based on within-species studies documenting higher survival of offspring located farther from conspecific adults and/or closer to light gaps. We determined whether the paradigm also holds among species by comparing spatial patterns of offspring mortality among three sympatric Neotropical rainforest tree species with the same seed dispersers but with different dispersal abilities. First, we assessed spatially non-random mortality for each species by measuring spatial shifts of the population recruitment curve (PRC) with respect to conspecific adults and light gaps across three early life stages: dispersed seeds, young seedlings and old seedlings. Then, we determined whether PRC shifts were greater for the species with short dispersal distances than for the species with greater dispersal distances. We found that the PRC shifted away from conspecific adults consistently across life stages, but we found no consistent PRC shifts towards gaps, suggesting that mortality was related more to the proximity of conspecifics than to absence of light gaps. PRC shifts away from adults were greatest in the species with the lowest dispersal ability, supporting the paradigm. Differential PRC shifts caused the spatial distribution of offspring to become almost independent with respect to adult trees and gaps in all three species, despite large differences in seed dispersal distance among these species. Our results provide direct empirical support for the paradigm that among tropical trees, species with farther seed dispersal are less impacted by spatially non-random mortality than are similar species with shorter dispersal distances. Thus, greater dispersal effectiveness merits extra investments of trees in seed dispersal ability, even at the cost of fecundity.     

Spatial and temporal patterns of seed attack and germination in a large-seeded neotropical tree species

It has long been argued that seed dispersal enhances recruitment in tropical trees by allowing offspring to `escape' strong density/distance-dependent attack by insects, pathogens and rodents. Here we examined the effects of canopy openness and parent-offspring distance upon the frequency and timing of Chlorocardium rodiei seed attack and germination within a 15-ha plot of Guyanan tropical rain forest. Seeds were artificially dispersed beneath parent trees, in the understorey away from trees and in gaps. Analysing our data from an 85-week period of regular monitoring, we found that the main spatial gradients, canopy openness and distance to nearest adult conspecific, do not lead to differences in the final number of seeds attacked by infesting scolytid beetles or rodents. The timing of beetle attack, however, varied along the distance gradient and this difference affords seeds at further distances a `window' in which to germinate and produce a seedling before attack. Canopy openness was not a good predictor of rooting success, but distance was strongly associated with root and shoot formation success and the mean time to shoot formation. There was a strong negative effect of distance on the likelihood of a seed being colonised by scolytid beetles prior to removal by rodents and shoot failure was strongly associated with prior infestation. We believe these results bring a key point to bear on the well-established notion of distance-dependent attack on seeds in tropical rainforests, viz. that seed characteristics (size, germination syndrome) and the timing of attack may be more important in explaining patterns of early seedling recruitment than distance. Our studies suggest that advantages accrued through dispersal in species like Chlorocardium will depend heavily on the `race' between seed germination and attack. In the case of Chlorocardium, the `race' can be lost at considerable distances due to its prolonged dormancy and the temporal fluctuations in fruitfall and rainfall which influence attack and germination. The results presented here suggest that the lag between seed attack and germination in tropical trees can regulate the influence of parent-offspring distance on cohort recruitment at this life history stage. Received: 5 March 1998 / Accepted: 6 December 1998     

Directed vertebrate-mediated seed dispersal of a fleshy-fruited amaryllid in a heterogenous habitat

Most plants with fleshy fruits have seeds that are ingested by animals, but a less well-understood mode of seed dispersal involves fleshy fruits containing seeds that are discarded by frugivorous animals because they are too large or toxic to be ingested. We studied the seed dispersal biology of Haemanthus deformis, an amaryllid lily species found in a mosaic of bush clumps in a grassland matrix in South Africa. We asked whether seed dispersal is directed in and among bush clumps and whether germination and survival are greater for seeds dispersed to bush clumps than for those dispersed into grassland. Using camera trapping, we found that fruits are consumed mainly by birds and rodents. The pulp was removed from the seeds which were then discarded without ingestion. While many seeds were dispersed close to the parent plant, most (c. 78.5%) were dispersed further than 1 m away from the parent plant. Longer distance dispersal resulted mainly from birds flying off with fruits in their bill or from rodents engaging in scatter-hoarding behavior. Seedling survival was most successful within bush clumps as compared to grasslands and shade was identified as a primary requirement for seedling survival. Seeds from which the fruit pulp had been removed germinated faster than those in intact fruits. Haemanthus deformis deploys a system of directed seed dispersal, whereby both birds and rodents contribute to the dispersal of seeds within patchy bush clumps that are favorable for seedling survival.     

Offspring spatial patterns in Picconia excelsa (Oleaceae) in the Canarian laurel forest

We studied the spatial patterns of seedlings and seeds in isolated Picconia excelsa (Oleaceae) trees in the laurel forest of Anaga, Tenerife (Canary Islands). By finding isolated trees we assessed the correlation of seed and seedling bank traits and parent trees by removing the confounding effects of proximity (<100 m radius) of conspecific fruiting trees. We counted all the seedlings per age (height) class within its parental range, and sampled the seed number along transects departing from beneath the parent canopy at regular intervals. We mapped all seedlings per age class and plotted seed and seedling profiles in relation to distance to parent trees. Older Picconia seedlings tended to clump significantly further from parent trees than younger seedlings, which clumped just beneath the parents. We found significant differences among distances to parent tree in numbers of seedlings per age class. The seedling bank area was significantly correlated with maximum distance of seedlings to parent trees. The majority of seeds were deposited within the first 4 m below the parent crown. Seedlings amount at further distances from the trees is larger than seeds/fruits as counted on the ground. Our results suggest that disseminated, older seedlings have occupied germination sites far from the parent tree because there is probably lower seedling–seedling and parent–seedling competition for resources, and perhaps no intraspecific allelopathy and predation/disease.     

Cheek-pouch dispersal of seeds by Japanese monkeys (Macaca fuscata yakui) on Yakushima Island, Japan

Seed dispersal by Japanese monkeys (Macaca fuscata yakui) via cheek-pouch was studied in a warm temperate evergreen forest on Yakushima Island. Plant list was compiled based on a study during 1986–1995, of which troops of monkeys have been habituated without artificial feeding. We followed the well-habituated monkeys in 1993 and 1994 to observe the feeding behavior and their treatments of fruits and seeds, and collected seeds dispersed by monkeys to record the distance carried from the mother trees. We checked the difference of germination ratio between seeds dispersed via cheek-pouch and seeds taken from mother trees by sowing experiments. Seeds and acorns of 22 species were observed to be dispersed via cheek-pouch of monkeys. Among them, three species with acorns were never dispersed via feces, and 15 species with drupes were seldom dispersed via feces. Plant species of which seeds are dispersed only via cheek-pouch had larger seeds than those of dispersed both via cheek-pouch and via feces, and typically had only one or two seeds in a fruit. As for one of cheek-pouch dispersal species,Persea thunbergii, the mean distance when seeds were carried from the mother trees via cheek-pouch was 19.7 m, and the maximum distance was as long as 105 m although more than 80% of seeds were dispersed within 30 m from mother trees. And 82% of seeds dispersed via cheek-pouch germinated. The easy separation of seeds from other parts of the fruit seems to facilitate cheek-pouch dispersal more than dispersal via feces. Cheek-pouch dispersal by monkeys has possibly enhanced the natural selection for larger seeds which bring forth larger seedlings with high shade-tolerance. In conclusion, cheek-pouch dispersal by monkeys is quite an important mode for trees in the mature stand in a warm temperate evergreen forest on Yakushima Island.     

Japanese maple (Acer palmatum var. Matsumurae, Aceraceae) recruitment patterns: seeds, seedlings, and saplings in relation to conspecific adult neighbors

We analyzed the spatial patterns among seeds, seedlings, saplings, and conspecific adult trees of the cool-temperate tree species Acer palmatum var. Matsumurae in a conifer-hardwood mixed forest in northern Japan, using two models that consider the influence of each adult within the neighborhood of the offspring. The results showed that recruitment patterns of each stage could be characterized and that significant shifts occur between successive stages. Sound seeds were more widely dispersed than unsound seeds; the mean dispersal distance (MDD) was 41.5 m for sound seeds, but only 12.6 m for unsound seeds. Most seedlings were located near conspecific adult trees, with a MDD of 14.3 m. Saplings, however, were more dispersed away from conspecific adult trees, with an MDD of more than 35 m. Light and gap distributions did not strongly affect the spatial distribution of the offspring; most saplings were located under nonconspecific canopies. These results suggest that the recruitment pattern of Japanese maple offspring is strongly affected by conspecific adult neighbors, rather than by light and gap distributions, with close proximity to conspecific adult trees reducing the growth and survival of seedlings during the transition to saplings.     

Factors Limiting Post‐logging Seedling Regeneration by Big‐leaf Mahogany (Swietenia macrophylla) in Southeastern Amazonia,Brazil, and Implications for Sustainable Management1

Post‐logging seedling regeneration density by big‐leaf mahogany (Swietenia macrophylla), a nonpioneer light‐demanding timber species, is generally reported to be low to nonexistent. To investigate factors limiting seedling density following logging within the study region, we quantified seed production rates, germinability, dispersal patterns, and seed fates on the forest floor through germination and the first seedling growing season in southeastern Amazonia, Brazil. Fruit production rates were low by three logged and one unlogged populations compared to reports from other regions. Commercial‐sized trees (>60 cm diameter) were more fecund than noncommercial trees (30–60 cm diameter) at two sites, averaging 14.5 vs. 3.9 fruits/tree/year, respectively, at Marajoara, a logged site, over 8 yr. Fruit capsules contained an average of 60.3 seeds/fruit, 70 percent of which appeared viable by visual inspection. Sixty‐seven to 72 percent of apparently viable seeds germinated in nursery beds 2.5 mo after the dispersal period, when wet season rains began. Dry season winds blew most seeds west‐northwest of parent trees, with median dispersal distances of 28 and 9 m on west and east sides of parent trees, respectively. Nearly 100 percent of seeds fell within an area of 0.91 ha. On the forest floor beneath closed canopies, mammals, invertebrates, and fungal pathogens killed 40 percent of apparently viable seeds, while 36 percent germinated. Nine months after seedling establishment—midway through the first logging season following seed dispersal—14 percent of outplanted seeds survived as seedlings, representing 5.8 seeds/fruit. We conclude that seedlings are likely to survive in logging gaps at appreciable densities only in rare cases where previous year fruit production rates by logged trees were high (4–12.5% of commercial‐sized trees/year at Marajoara) and where tree crowns were felled in west or northwest directions.     

Seed and seedling ecology of piñon and juniper species in the pygmy woodlands of western North America

Knowledge of the seed and seedling ecology of the piñon and juniper woodlands of western North America is essential for understanding both the northward migration and expansion of the woodlands during the Holocene (< 11,500 B.P.), and the accelerated expansion of the woodlands since settlement of the West by Anglo-Americans around 200 years ago. We follow the fates of seeds and seedlings of the different piñon and juniper species within the woodlands from seed development to seedling establishment, and discuss the implications of this information for the past and present expansion of the woodlands. While seed development requires about two and one-half years in pinons, it is species-dependent in junipers and can take one, two, or even three years. Substantial seed losses can occur during seed development due to developmental constraints, and before or after seed maturation as a result of insects, pathogens, or predatory animals. In piñon pines, the primary seed dispersers are scatterhoarding birds (corvids) and rodents that harvest seeds from the trees or after seed fall and cache them in the soil. In contrast, most junipers appear to be dispersed primarily by frugivorous birds and mammals that ingest the seeds and defecate them onto the soil surface. We have recently documented that scatter-hoarding rodents also disperse juniper seeds. Disperser effectiveness, or the contribution a disperser makes to the future reproduction of a plant population, may vary among species of piñons and especially junipers. Piñon seeds are short-lived and exhibit little dormancy, and they probably only germinate the spring following dispersal. Juniper seeds are long-lived and seed dispersal can occur over one or more years. Seed germination can be delayed for several years due to impermeable seed coats, embryo dormancy, or the presence of inhibitors. Seedling establishment of piñon pines is facilitated by nurse plants but, while junipers often establish beneath nurse plants, they are capable of establishing in open environments. In the southwestern United States, higher establishment of juniper occurs in open environments due to more favorable precipitation, and competition may be more important than facilitation in determining establishment. When considering the mechanisms involved in the past and present expansion of the woodlands, short-distance dispersal, local population growth, and long-distance dispersal are all important. Different classes of dispersers, some of which appear to have coevolved with the tree species, appear to be responsible for local (short-distance) vs. long-distance dispersal in pinons and junipers. Because ecotones form the interface between the woodlands and adjacent communities, they can provide valuable information on both the seed dispersal and seedling establishment processes responsible for tree expansion. Disturbance regimes and, recently, the effects of humans on those regimes have major effects on the expansion and contraction of the woodlands. Before Anglo-American settlement, fires occurred as frequently as every 50–100 years throughout much of the woodlands. During this century, fire frequencies have been reduced due to the indirect effects of livestock grazing and the direct effects of removing Native Americans from the ecosystem and implementing active fire-prevention programs. The result has been an increase in tree-dominated successional stages at the expense of grass-dominated stages. Various management techniques, including controlled burning and chaining, have been implemented to reduce tree dominance, but their effects depend largely on the life histories of the tree species and the disturbance characteristics. Several areas relating to the seed and seedling ecology of the piñon and juniper require additional research if we are to truly understand the dynamics of the woodlands.     

Seed caching by woylies Bettongia penicillata can increase sandalwood Santalum spicatum regeneration in Western Australia

Abstract The role a small marsupial, the woylie Bettongia penicillata, might play in the recruitment and regeneration of Western Australian sandalwood Santalum spicatum through its seed caching behaviour was investigated in this study. To determine the fate of the seeds, cotton thread was attached to the seeds and the trail followed. A total of 25 seed caches were located. All of the seeds were found in separate caches, which was consistent with scatter‐hoarding behaviour. The average distance from the source of the seeds to the cache was 43.1 m ± 5.8 m at Dryandra woodland and 29.1 m ± 3.8 m at Karakamia sanctuary. The mean cache depth was 4.3 cm ± 0.2 cm at Dryandra woodland compared with 4.6 cm ± 0.3 cm at Karakamia sanctuary. Significantly more seedlings and saplings grew away from sandalwood trees at sites where woylies were present than at sites with no woylies. In contrast, significantly more seedlings and saplings grew under adult sandalwood trees at the site without woylies than where they were present, although there were significantly lower rates of recruitment and sandalwood regeneration at these sites. In addition, significantly more whole, undisturbed sandalwood seeds were found under the parent trees at the woylie‐free site than at the site with woylies. These findings strongly suggest that little seed dispersal or regeneration of sandalwood occurs in the absence of woylies. Through scatter‐hoarding, woylies have the potential to disperse and cache sandalwood seeds away from the source and significantly alter the subsequent regeneration of sandalwood. Furthermore, by caching seeds large distances away from a source, woylies could modify the distribution of sandalwood in an area.     

A test of the escape and colonization hypotheses for zoochorous tree species in a Western Amazonian forest

In order to assess the importance of seed dispersal (escape and colonization hypotheses), I used transplant experiments for seeds and seedlings of 5–11 plant species with fleshy fruits in a lowland tropical forest (Tinigua National Park, Colombia). I controlled seed density, distance to parental tree, and habitat type. I monitored seed removal, seedling survival, and seedling growth during the first year of development for an average of 554 seeds and 169 seedlings for each species. I supplemented the experimental results with measurements of natural recruitment. I found little support for the escape hypothesis during the seed and seedling stages. For six species that showed differences in seed removal associated with distance, five showed highest removal away from, than close to parent trees, suggesting predator satiation. Seedling survival during the first year was not consistently associated with low densities and long distances from parent trees. For the majority of species, seedlings did not survive flooding in low basins, and there was growth advantage for most plant species in canopy gaps. These differences imply advantages for seed dispersal to adequate habitats, as predicted by the colonization hypothesis. In contrast to experiments, strong negative distance-dependent effects were evident when analyzing natural recruitment patterns. The ratio between saplings and seedlings was higher away from parent trees for the species with enough recruitment to be analyzed and this suggests that a negative distance-dependent effect may also occur after seedling establishment. This pattern is suspected for several other species, but an analysis with some of the other most common trees showed a variety of negative, neutral, and positive distance dependent effects. This study emphasizes the importance of long-term studies to asses the role of seed dispersal.     

Patterns of pollen and seed dispersal in a small, fragmented population of the wind-pollinated tree Araucaria angustifolia in southern Brazil

Paternity analysis based on eight microsatellite loci was used to investigate pollen and seed dispersal patterns of the dioecious wind-pollinated tree, Araucaria angustifolia. The study sites were a 5.4 ha isolated forest fragment and a small tree group situated 1.7 km away, located in Paranalpha State, Brazil. In the forest fragment, 121 males, 99 females, 66 seedlings and 92 juveniles were mapped and genotyped, together with 210 seeds. In the tree group, nine male and two female adults were mapped and genotyped, together with 20 seeds. Paternity analysis within the forest fragment indicated that at least 4% of the seeds, 3% of the seedlings and 7% of the juveniles were fertilized by pollen from trees in the adjacent group, and 6% of the seeds were fertilized by pollen from trees outside these stands. The average pollination distance within the forest fragment was 83 m; when the tree group was included the pollination distance was 2006 m. The average number of effective pollen donors was estimated as 12.6. Mother-trees within the fragment could be assigned to all seedlings and juveniles, suggesting an absence of seed immigration. The distance of seedlings and juveniles from their assigned mother-trees ranged from 0.35 to 291 m (with an average of 83 m). Significant spatial genetic structure among adult trees, seedlings, and juveniles was detected up to 50 m, indicating seed dispersal over a short distance. The effective pollination neighborhood ranged from 0.4 to 3.3 ha. The results suggest that seed dispersal is restricted but that there is long-distance pollen dispersal between the forest fragment and the tree group; thus, the two stands of trees are not isolated.     

Patterns of Seed Dispersal and Dispersal Failure in a Hawaiian Dry Forest Having Only Introduced Birds

Dry forests are among the most endangered natural communities in the Hawaiian Islands. Most have been reduced to isolated trees and small forest fragments in which native tree species reproduce poorly. The replacement of native birds by introduced generalists may be contributing to dry forest decline through modification of seed dispersal patterns. To document seed dispersal by introduced birds, we conducted foraging observations on fleshy-fruited trees and measured seed rain under trees and in adjacent open areas for 1 year in a dry forest dominated by native trees. Although trees covered only 15.2 percent of the study area, 96.9 percent of the bird-dispersed seeds were deposited beneath them. The Japanese white-eye (Zosterops japonicus) was the principal dispersal agent. Among bird-dispersed seeds, those of the invasive tree Bocconia frutescens accounted for 75 percent of all seeds collected beneath trees (14.8 seeds/m²/yr) and the invasive shrub Lantana camara accounted for 17 percent. Although nearly 60 percent of the reserve's native woody species possess fleshy fruits, introduced birds rarely disperse their seeds. Native trees accounted for <8 percent of all bird-dispersed seeds and are consequently experiencing dispersal failure by falling directly under parent trees. Smaller-seeded non-native plants, in contrast, may be benefiting from dispersal by introduced birds. Current dispersal patterns suggest that these readily disseminated non-native plants may eventually replace the remaining native flora.     

Dispersal of Eucalyptus torelliana seeds by the resin-collecting stingless bee,Trigona carbonaria

Ants are the only group of invertebrates currently identified as significant dispersers of seeds, but we report here the dispersal of Eucalyptus torelliana seeds by bees. Fruits of E. torelliana produce resin which is collected by workers of the stingless bee Trigona carbonaria. Seeds adhere to resin in the workers' corbiculate and are transported to the nest. Workers transported seeds distances of more than 300 m from the parent tree and seeds at the nest were viable and capable of germination. Seeds were removed from the nests by workers and discarded away from the nest, and E. torelliana trees became established in the vicinity of colonies of T. carbonaria. Mellitochory may be a novel method of seed dispersal where bees are dispersers, and associated with fruits that produce resin as an attractant for bees.     

R.M. Cowling,D.M. Richardson & S.M. Pierce (eds.) 1977. Vegetation of Southern Africa.

We determined the role of bird dispersal in seed and seedling dynamics of the tree Kalopanax pictus from 7 years of observing seed rain and seedling emergence in a broad-leaved deciduous forest in central Japan. We also performed an experiment on the influence of seed pulp on germination of seeds of K. pictus. Seeds of this species can lie dormant for several years, and this causes rather constant yearly seedling emergence in spite of irregular seed production. The spatial distribution of the seedlings that emerged each year (maximum distance from nearest conspecific seed-bearing tree of 90 m) was wider than that of gravity-dispersed seeds (max. distance of 37 m), suggesting seed dispersal by birds in winter. Emerged seedling densities at sites over 20 m from the nearest conspecific seed-bearing tree were highest in the spring of 1991, about half a year after the largest seed fall of the observation period. However, emerged seedling densities within 20 m from seed-bearing trees were highest in 1992, 1.5 years after the largest seed fall. These field observations may be explained by the experimental results on the effects of seed pulp on germination. Intact seeds germinate slowly at low germination rates, while seeds without seed pulp germinate quickly at high germination rates. Fallen seeds with seed pulp thus appear to form a seed bank near seed sources (temporal dispersal), while seeds scattered by birds appear to increase the possibility of reaching the present safe sites in distant areas with quick germination (spatial dispersal).