Fruit removal and postdispersal survivorship in the tropical dry forest shrub Erythroxylum havanense: ecological and evolutionary implications

We studied the relationship between the removal rate and the spatiotemporal availability of ripe fruits of the tropical deciduous shrub Erythroxylum havanense in western Mexico. We also evaluated the effects of dispersal on seed survival during the first stages of establishment. Fast and early dispersal should be favored in E. havanense, since propagules have more time to grow and accumulate resources before the beginning of the severe dry season. In general, high rates of fruit removal imply faster and earlier dispersal. Thus, plants producing large crops should benefit from high removal rates, which will increase the probability of successful establishment by their progeny. To characterize both individual and population fruiting patterns, we made daily counts of fruits on 51 plants arranged in six clumps of different sizes. The daily number of fruits removed per plant was higher for plants with larger initial crop sizes and larger numbers of ripe fruits on a given day, but decreased as clump size increased. Additionally, we monitored postdispersal survival and germination in an experiment manipulating seed density, distance from adult plants, and seed predation. Early establishment was independent of density or distance, and vertebrate seed predation was the main agent of seed mortality. Our results indicate that the critical variable with respect to fruit removal is the number of fruits a plant produces, large plants having higher dispersal rates. Large plants are also more likely to have more seeds escaping postdispersal seed predation.     

Fruit removal efficiency and success: influence of crop size in a neotropical treelet

The reproductive success of animal-dispersed plants is closely linked to the number of seeds that they are able to disperse. The fruit crop size hypothesis states that a plant with large fruit crop size will attract more dispersers than a plant with a smaller fruit crop, which may result in more seeds being dispersed from the foremost. In this study, we experimentally examined the effect of crop size and other factors on primary seed dispersal in a neotropical shrub/tree, Casearia corymbosa (Flacourtiaceae). We used two predictive variables of reproductive success, which produce an accurate picture of seed dispersal ratio: fruit removal efficiency (proportion of a fruit crop removed by frugivores) and fruit removal success (relative contribution of each individual tree to the number of fruits removed in the population). We established two levels of fruit crop size at the C. corymbosa individuals, using plants with large (150 fruits) and small crops (50 fruits). We found that individual plants with larger crops had significantly higher values of fruit removal efficiency (92.6%) and success (5%) than plants with smaller crops (69.3% and 1.3%, respectively). Fruit removal efficiency was related to vegetation type, plant height and fruit width, but the variance explained by these variables was low ( < 8%). Fruit removal success was significantly related to crop size ( > 90% of the variance explained). These results suggest that fruit removal efficiency and success are strongly related to fruit crop size of C. corymbosa plants.     

Seed size selection by olive baboons

Seed size is an important plant fitness trait that can influence several steps between fruiting and the establishment of a plant’s offspring. Seed size varies considerably within many plant species, yet the relevance of the trait for intra-specific fruit choice by primates has received little attention. Primates may select certain seed sizes within a species for a number of reasons, e.g. to decrease indigestible seed load or increase pulp intake per fruit. Olive baboons (Papio anubis, Cercopithecidae) are known to select seed size in unripe and mature pods of Parkia biglobosa (Mimosaceae) differentially, so that pods with small seeds, and an intermediate seed number, contribute most to dispersal by baboons. We tested whether olive baboons likewise select for smaller ripe seeds within each of nine additional fruit species whose fruit pulp baboons commonly consume, and for larger seeds in one species in which baboons feed on the seeds. Species differed in fruit type and seed number per fruit. For five of these species, baboons dispersed seeds that were significantly smaller than seeds extracted manually from randomly collected fresh fruits. In contrast, for three species, baboons swallowed seeds that were significantly longer and/or wider than seeds from fresh fruits. In two species, sizes of ingested seeds and seeds from fresh fruits did not differ significantly. Baboons frequently spat out seeds of Drypetes floribunda (Euphorbiaceae) but not those of other plant species having seeds of equal size. Oral processing of D. floribunda seeds depended on seed size: seeds that were spat out were significantly larger and swallowed seeds smaller, than seeds from randomly collected fresh fruits. We argue that seed size selection in baboons is influenced, among other traits, by the amount of pulp rewarded per fruit relative to seed load, which is likely to vary with fruit and seed shape.     

The effect of frugivory on postdispersal seed removal and germination in the pantropical forest tree Antiaris toxicaria Leschenault

The quality of seed treatment by frugivores has an effect on seed removal after dispersal, seed germination and tree recruitment. We provide information on postdispersal seed removal, germination and subsequent recruitment in tropical forest tree species Antiaris toxicaria in Ghana. We tested whether postdispersal seed removal and germination rates were differentially affected by the following seed treatments: seeds that were spat out by monkeys with all fruit pulp removed and spitting seeds with fruit pulp partially removed as observed in some birds and bats. We used seeds of intact ripened fruits as control. Frugivore seed treatment and distance from bole affected seed removal patterns, whereas intact seeds were significantly removed from all seed stations. The germination success was greater for seeds that were spat out by monkeys and poor for seeds with fruit pulp partially removed and intact fruits. More recruits were recorded at the edge of the adult A. toxicaria canopy radius. There was weak relationship (r² = 0.042) between the number of recruits and distance away from the adult tree. Results suggest that the subsequent recruitment in tropical forest tree species may be enhanced by some frugivore fruit‐handling behaviour where fruit pulp is removed from the seeds without destroying the seeds.     

Satiation of predispersal seed predators: the importance of considering both plant and seed levels

Plants can reduce the fitness costs of granivory by satiating seed predators. The most common satiation mechanism is the production of large crops, which ensures that a proportion of the seeds survive predation. Nevertheless, satiation of small granivores at the seed level may also exist. Larger seeds would satiate more efficiently, enhancing the probability of seed survival after having been attacked. However, a larger seed size could compromise the efficiency of satiation by means of large crops if there were a negative relationship between seed size and the number of seeds produced by an individual plant. We analyze both types of satiation in the interaction between the holm oak Quercus ilex and the chestnut weevil Curculio elephas. Both crop size and acorn size differed strongly in a sample of 32 trees. Larger crop sizes satiated weevils, and higher proportions of the seeds were not attacked as crop size increased. Larger seeds also satiated weevil larvae, as a larger acorn size increased the likelihood of embryo survival. Seedling size was strongly related to acorn size and was reduced by weevil attack, but seedlings coming from large weeviled acorns were still larger. The number and the size of the acorns produced by individual trees were negatively related. Larger proportions of the crop were infested in oaks producing less numerous crops of larger acorns. However, contrary to expectations, these trees did not satiate more effectively at the seed level either. Effective satiation by larger acorns was precluded by larger multi-infestation rates associated to smaller seed crops, in such a way that the proportion of attacked seeds that survived did not vary among trees with different acorn sizes. These results highlight the need of considering satiation by means of large crops and large seeds in studies of predispersal seed predation. Long-term monitoring on individual oaks will help to assess whether there is a trade-off between the number and the size of the acorns and, if it existed, how it could condition the fitness consequences of both types of satiation.     

Pre‐dispersal seed predators and fungi differ in their effect on Luehea seemannii capsule development,seed germination,and dormancy across two Panamanian forests

Pre‐dispersal seed predation can greatly reduce crop size affecting recruitment success. In addition, non‐fatal damage by seed predators may allow infection by fungi responsible for post‐dispersal seed losses. The objectives of this study were (1) to quantify pre‐dispersal seed predation and fungal infection in a Neotropical tree species, Luehea seemannii, that produces dehiscent fruits and wind‐dispersed seeds, and (2) to link pre‐dispersal effects on seed quality to seed survival in the soil. To examine how seed predators and fungi influence seed losses, mesh exclosures, fungicide, and the combination of both treatments were applied to separate branches in the canopy of trees in Gamboa and Parque Natural Metropolitano (PNM), Panama. To determine if treatments affect seed viability and survival in the soil, half of the seeds collected from each treatment were buried for 4 weeks in forest soils and subsequently allowed to germinate before and after the breaking of dormancy. Overall, 24 percent of developing fruit were lost to insect attack. In contrast, fungi infected only 3 percent of seeds at the pre‐dispersal stage. For seeds germinated directly after collection, fungicide significantly increased germination in the wetter site (Gamboa) but decreased germination in the drier site (PNM). The pre‐dispersal insect exclosure treatment increased the fraction of seeds that remained dormant after burial in the soil. This result suggests that exposure to insect predators may cause physical damage to seeds that results in the loss of physical dormancy but does not necessarily increase the susceptibility of seeds to pathogen attack in the soil.     

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.     

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     

Spatial dynamics of <Emphasis Type="Italic">Ilex aquifolium</Emphasis> populations seed dispersal and seed bank: understanding the first steps of regeneration

The objective of this study was to analyse quantitatively the spatial distribution of holly (Ilex aquifolium L.) seed rain and seed bank, and to detect the relationships between these consecutive processes. We measured seed dispersal by birds and fallen fruits, and also density and viability of seed bank in two Ilex populations in central Spain. Analysis was made distinguishing the following microhabitats: holly woodland, edge of holly woodland, open grassland 10 m and 100 m from the woodland, fleshy fruit shrubs, dry fruit shrubs, and adjacent non-holly woodland. Spatial distribution of dispersed and in-soil seeds was measured by the clumping index. Seed rain and seed bank under holly woodlands were significantly higher than in the other microhabitats. Forest edges and fleshy fruit shrubs were the next microhabitats with the highest seed rain and seed bank density. Interannual and interlocality variations were not significant. The relative importance of the different dispersal methods varied between microhabitats, with a similar support of bird dispersed seeds and fallen fruits within the woodland and a greater influence of cattle dispersal in open areas. Seed spatial aggregation was significant in both dispersed seeds and soil seeds from holly woodlands and the edge of the forest. Aggregation under shrubs, grasslands and the adjacent forests evidenced a general random distribution of holly seeds (only in some cases clumping index was significant). Quantitative differences between seed rain and seed bank are important. Post-dispersal seed predation did not modify seed rain distribution, which was mirrored in the seed bank pattern. These two phases of holly regeneration had a heavy spatial influence, determined by the landscape structure and activity of the dispersal agents, that reflects a differential recruitment potential. Comparisons between both populations suggest that in the southern locality (Robregordo) holly has a weaker capacity to colonize open areas, and a stronger recruitment limitation due to propagule availability.     

Evolution of a unique seed maturity pattern in Croton bonplandianum Baill strengthens ant-plant mutualism for seed dispersal

Summary The female flowers of Croton bonplandianum bear nectar glands which become active during fruit maturation and attain peak activity just prior to the splitting of fruits. This temporal specificity of nectar gland activity is shown to facilitate seed dispersal by ants, which are attracted to the plant only during the fruit maturation period. The nectar glands establish a nectar influence zone with a radius of 60 cm around the plant within which seed dispersal by ants is effective. Seed dispersal by ants is facilitated only if the seeds are placed within this nectar influence zone. This is accomplished by an intriguing evolutionary shift in the maturation pattern of the fruits. Unlike the usual acropetal development, fruit maturation in Croton is temporally asymmetrical, with the fruits nearer the parental axis maturing early. This unique pattern of fruit development together with the polychasial branching system leads to a concentration of seeds within the nectar influence zone and enhances seed dispersal by ants. The proximate factors responsible for this asynchronous fruit maturity were investigated.     

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.     

Responses of dispersal agents to tree and fruit traits in Virola calophylla (Myristicaceae): implications for selection

Variation in traits affecting seed dispersal in plants has been attributed to selection exerted by dispersal agents. The potential for such selection was investigated in Virola calophylla (Myristicaceae) in Manú National Park, Peru, through identification of seed dispersal agents and of tree and fruit traits significantly affecting the quantity of seeds dispersed. Seventeen bird and one primate species (the spider monkey, Ateles paniscus) dispersed its seeds. Spider monkeys dispersed the majority of seeds (a minimum of 83% of all dispersed seeds). Visitation by dispersal agents depended only on the quantity of ripe fruit available during a tree observation. In contrast, seed removal increased with both greater quantity of ripe fruit and aril: seed ratio. When analyzed separately, seed removal by birds increased only with greater aril: seed ratio, whereas seed removal by spider monkeys was affected by the quantity of ripe fruit and phenological stage. The finding that dispersal agents responded differently to some tree and fruit traits indicates not only that dispersal agents can exert selection on traits affecting seed dispersal, but also that the resulting selection pressures are likely to be inconsistent. This conclusion is supported by the result that the proportion of the seed crop that was dispersed from individual trees, which accounted for cumulative dispersal by all agents, was not influenced by any tree or fruit trait evaluated. Comparing these results with those from studies of V. sebifera and V. nobilis in Panama revealed that the disperser assemblages of these three Virola species were congruent in their similar taxonomic representation. In Panama the proportion of V. nobilis seed crop dispersed was related positively to aril: seed ratio and negatively to seed mass, a result not found for V. calophylla in Peru. The greater importance of dispersal by primates versus birds in V. calophylla, relative to V. nobilis, may explain this difference. Thus, variation in disperser assemblages at regional scales can be another factor contributing to inconsistency in disperser-mediated selection on plant traits.     

Effects of seed size on dispersal distance in five rodent-dispersed fagaceous species

We studied the effect of seed size on dispersal by comparing dispersal distances in five rodent-dispersed fagaceous species (Lithocarpus harlandii, Quercus variabilis, Q. serrata, Cyclobalanopsis glauca, Castanopsis fargesii) with different seed size. We tracked individual seeds with coded tin-tags in two stands over 3 years in a subtropical evergreen broadleaved forest in the Dujiangyan Region of Sichuan Province, Southwest China. Our seed tracking data indicate that dispersal distances (including mean, maximum and distribution range) of seeds in primary caches and of seeds eaten after dispersal significantly increased with seed size, for both stands and all years. In addition, larger seeds (L. harlandii and Q. variabilis) were re-cached more often than smaller ones, which further reduced the relative density among caches and extended dispersal distances. Our findings indicate that greater dispersal distances for larger seeds might benefit the evolution of differences in seed size, and that scatter-hoarding might be advantageous for rodent-dispersed tree species.     

Effect of distance,aggregation, and habitat on levels of seed predation for two mammal — dispersed neotropical rain forest tree species

The effect of seed aggregation and distance from conspecific trees on seed predation was experimentally examined for two neotropical tree species, Macoubea guianensis (Apocynaceae) and Pouteria sp. (Sapotaceae) in a lowland tropical rain forest in northeastern Peru. Results of these experiments are discussed in the context of the Janzen-Connell model (Janzen 1970; Connell 1971), which predicts decreased seed survival near parent trees due to either density-or distance-responsive mortality, and Howe's model (Howe 1989) which predicts that trees with seeds dispersed in clumps (aggregated) will not suffer density-dependent predation, and will have higher survival of seeds near the parent tree than other trees. We also examined whether predation on seeds of these species was affected by seed placement in or near 30-m-wide strips regenerating after clear-cutting. Both species appeared to be mammal-dispersed but differed in how frugivores handled seeds, seed size, overall fruit crop size, and gemination time. Neither of the two species studied appeared to suffer seed predation in a manner predicted by the Janzen-Connell model, and patterns of seed predation for only one of the species was similar to predictions of Howe's model. For neither species did seed predation along the edge of, or in the center of, regenerating clear cuts differ from predation 15 m into the primary forest. For Pouteria, seed predation in and near regnerating strips was significantly greater than around forest trees, but the opposite pattern held for Macoubea. Overall, seed predation was much greater on Macoubea. The difference in seed predation for these two species was most likely a result of differences in the types of seed predators that attacked these two species.     

应用结构方程模型解析影响黄连木果实产量 和种子命运的因素

果实(种子)产量和质量是影响植物种群更新的重要因素。为了探明影响黄连木果实产量和种子命运的因素以及这些影响因素之间的相互作用, 作者于2009年对河南省济源市45株黄连木(Pistacia chinensis)结果样树的植株特征、果实特征、果实产量和种子命运等进行了测定, 并用结构方程模型进行综合分析。结果表明: (1)黄连木果实产量与树高、树冠面积和果序大小等特征成正相关, 但与胸径、果实大小相关性不显著; (2)与捕食者饱和假说的预测不一致, 单株果实产量对黄连木广肩小蜂(Eurytoma plotnikovi)的种子捕食率(即虫蛀率)无显著直接负向效应; (3)树高和果实大小对虫蛀率为显著直接正向效应, 胸径对虫蛀率为显著直接负向效应, 显示黄连木广肩小蜂对植株特征和果实特征有一定的选择能力; (4)空壳率与虫蛀率成显著负相关, 空壳果实越多, 越易逃避黄连木广肩小蜂的寄生, 空壳果实的存在对完好种子起到了一定保护作用, 可能是黄连木防御昆虫寄生的重要机制; (5)空壳率和虫蛀率对种子完好率有显著直接负向效应, 而胸径、果序大小和果实产量对完好率为间接正向效应, 树高和果实大小为间接负向效应。可见, 黄连木植株特征和果实特征均不同程度地影响其果实产量和昆虫寄生, 从而影响黄连木的种子质量和种群更新。     

Seed dynamics of resprouting shrubs in grassy woodlands: Seed rain,predators and seed loss constrain recruitment potential

Abstract Measuring the fate of seeds between seed production and seedling establishment is critical in understanding mechanisms of recruitment limitation of plants. We examined seed fates to better understand the recruitment dynamics of four resprouting shrubs from two families (Fabaceae and Epacridaceae) in temperate grassy woodlands. We tested whether: (i) pre‐dispersal seed predation affected seed rain; (ii) post‐dispersal seed predation limited seed bank accumulation; (iii) the size of the seed bank was related to seed size; and (iv) viable seeds accumulated in the soil after seed rain. There was a distinct difference in seed production per plant between plant families with the legumes producing significantly more seeds per individual than the epacrids. Seed viability ranged from 43% to 81% and all viable had seed or fruit coat dormancy broken by heat or scarification. Pre‐dispersal predation by Lepidopteran larvae removed a large proportion of seed from the legume seed rain but not the epacrids. Four species of ants (Notoncus ectatomoides, Pheidole sp., Rhytidoponera tasmaniensis and Iridomyrmex purpureus) were major post‐dispersal seed removers. Overall, a greater percentage of Hardenbergia (38%) and Pultenaea (59%) seeds were removed than the fleshy fruits of Lissanthe (14%) or Melichrus (0%). Seed bank sizes were small (<15 seeds m^?2) relative to the seed rain and no significant accumulation of seed in the soil was detected. Lack of accumulation was attributed to seed predation as seed decay was considered unlikely and no seed germination was observed in our study sites. Our study suggests that seed predation is a key factor contributing to seed‐limited recruitment in grassy woodland shrubs by reducing the number of seeds stored in the soil.     

Crop size,plant aggregation,and microhabitat type affect fruit removal by birds from individual melastome plants in the Upper Amazon

We studied the efficiency (proportion of the crop removed) and quantitative effectiveness (number of fruits removed) of dispersal of Miconia fosteri and M. serrulata (Melastomataceae) seeds by birds in lowland tropical wet forest of Ecuador. Specifically, we examined variation in fruit removal in order to reveal the spatial scale at which crop size influences seed dispersal outcome of individual plants, and to evaluate how the effect of crop size on plant dispersal success may be affected by conspecific fruit abundance and by the spatial distribution of frugivore abundance. We established two 9-ha plots in undisturbed terra-firme understory, where six manakin species (Pipridae) disperse most seeds of these two plant species. Mean levels of fruit removal were low for both species, with high variability among plants. In general, plants with larger crop sizes experienced greater efficiency and effectiveness of fruit removal than plants with smaller crops. Fruit removal, however, was also influenced by microhabitat, such as local topography and local neighborhood. Fruit-rich and disperser-rich patches overlapped spatially for M. fosteri but not M. serrulata, nonetheless fruit removal of M. serrulata was still much greater in fruit-rich patches. Fruit removal from individual plants did not decrease in patches with many fruiting conspecifics and, in fact, removal effectiveness was enhanced for M. fosteri with small crop sizes when such plants were in patches with more conspecifics. These results suggest that benefits of attracting dispersers to a patch balanced or outweighed the costs of competition for dispersers. Spatial pattern of fruit removal, a measure of plant fitness, depended on a complex interaction among plant traits, spatial patterns of plant distribution, and disperser behavior.     

Ficus seed shadows in a Bornean rain forest

Due to their copious seed production and numerous dispersers, rain forest fig trees have been assumed to produce extensive and dense seed shadows. To test this idea, patterns of seed dispersal of two species of large hemiepiphytic fig tree were measured in a Bornean rain forest. The sample included four Ficus stupenda and three F. subtecta trees with crop sizes ranging from 2,000 to 40,000 figs (400,000 to 13,000,000 seeds). Seed rain out to a distance of 60 m from each study tree was quantified using arrays of seed traps deployed in the understory. These trees showed a strongly leptokurtic pattern of dispersal, as expected, but all individuals had measurable seed rain at 60 m, ranging from 0.2 to 5.0 seeds/m². A regression of In-transformed seed rain density against distance gave a significant fit to all seven trees' dispersal patterns, indicating that the data could be fitted to the negative exponential distribution most commonly fitted to seed shadows. However, for six of seven trees, an improved fit was obtained for regressions in which distance was also In-transformed. This transformation corresponds to an inverse power distribution, indicating that for vertebrate-dispersed Ficus seeds, the tail of the seed rain distribution does not drop off as rapidly as in the exponential distribution typically associated with wind dispersed seed shadows. Over 50% of the seed crop was estimated to fall below each fig tree's crown. Up to 22% of the seed crop was dispersed beyond the crown edge, but within 60 m of the tree. Estimates of the maximum numbers of seeds which could have been transported beyond 60 m were 45% for the two largest crops of figs, but were under 24% for the trees with smaller crops. Seed traps positioned where they had an upper canopy layer above them were associated with higher probabilities of being hit by seeds, suggesting that vertebrate dispersal agents are likely to perch or travel through forest layers at the same level as the fig crown and could concentrate seeds in such areas to some degree. The probability of a safe site at 60 m from the fig tree being hit by seeds is calculated to be on the order of 0.01 per fruiting episode. Fig trees do not appear to saturate safe sites with seeds despite their large seed crops. If we in addition consider the rarity of quality establishment sites and post-dispersal factors reducing successful seedling establishment, hemiepiphytic fig trees appear to face severe obstacles to seedling recruitment.     

Effects of harvester ants on seed availability and dispersal of Lavandula stoechas subsp. pedunculata in a Mediterranean grassland-scrubland mosaic

The role of harvester ants in Mediterranean grassland and scrubland has mostly focused on seed consumption. However, recent studies have reported their role as accidental dispersal agents of some of the collected seeds via refuse piles. The objective of this study is to examine the effect of the ant Messor barbarus on seed availability and dispersal of one of its major diet components, Lavandula stoechas subsp. pedunculata, in scrubland, grassland and the ecotones between them. After confirming and quantifying the Lavandula contribution to M. barbarus diet, we described the spatial and temporal patterns of pre- and post-dispersal seed predation, seed content and seedling occurrence in the refuse piles. Our results show that: (1) Lavandula propagules constitute a high proportion of the prey items collected by M. barbarus, with particularly intense collection activity in mid-summer, spring and autumn, in decreasing order. (2) Pre-dispersal predation rate was significantly higher in the ecotone than in the scrubland (76% and 13.5% of total seed production lost respectively). (3) Season and propagule type (seed vs. fruit) were the most significant variables explaining the post-dispersal predation probability, which approached 100% of seeds after 48 h in mid-summer. (4) Viable Lavandula seeds were found in refuse piles at densities of 0.06–0.2 per g of refuse pile material, or 58.8–207.2 per refuse pile. On the one hand, these results indicate that the ecotones are most affected by M. barbarus pre-dispersal consumption, which may locally limit Lavandula colonisation. On the other hand, the small proportion of consumed seeds that is dispersed to refuse piles may be relevant at the population level, as this dispersal implies arrival at potentially favourable sites for establishment.     

The role of Orii’s flying-fox (Pteropus dasymallus inopinatus) as a pollinator and a seed disperser on Okinawa-jima Island, the Ryukyu Archipelago, Japan

The role of the Orii’s flying-fox (Pteropus dasymallus inopinatus) as a pollinator and a seed disperser on Okinawa-jima Island was investigated by direct observations and radio-tracking from October 2001 until January 2006. We found that Orii’s flying-fox potentially pollinated seven native plant species. Its feeding behavior and plant morphological traits suggested that this species is an important pollinator of Schima wallichii liukiuensis and Mucuna macrocarpa. The flying-fox also dispersed the seeds of 20 native plant species. The seeds of all plants eaten by the flying-fox were usually dropped beneath the parent tree, although large fruits of four plant species were occasionally brought to the feeding roosts in the mouth, with the maximum dispersal distance—for Terminalia catappa—estimated to be 126 m. Small seeds of 11 species (mostly Ficus species) were dispersed around other trees, during the subsequent feeding session, through the digestive tracts, with the mean dispersal distance for ingested seeds estimated at 150 ± 230.3 m (±SD); the maximum dispersal distance was 1833 m. A comparison of the seed dispersal of available fruits according to the size of flying-foxes and other frugivores suggested that the seed dispersal of eight plant species producing large fruits mostly depended on Orii’s flying-fox. On Okinawa-jima Island, the Orii’s flying-fox plays an important role as a pollinator of two native plants and as a long-distance seed disperser of Ficus species, and it functions as a limited agent of seed dispersal for plants producing large fruits on Okinawa-jima Island.