天然次生林中水曲柳种子的扩散格局

 种子扩散是种群动态中的一个关键环节,研究其扩散格局对了解种子库及更新格局有重要意义。于1999～2000年在天然次生林中选择样地并设置样带,共设取样点1376个,观察和估计了水曲柳(Fraxinus mandshurica)的种子雨格局。结果发现,母树附近,尤其是冠幅范围内,各方向的种子雨没有显著差别。但种子雨密度随距离有明显变化,冠幅范围内平均可达462～903个·m-2。距冠缘10～20 m急剧减少,为50～190个·m-2。72.5%～96.4%的种子散落在距母株树干10 m 的范围内。饱满种子顺风     

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.     

Vertical stratification of figs and fig-eaters in a Bornean lowland rain forest: how is the canopy different?

Fig trees (Ficus spp; Moraceae) are a common constituent of many tropical forests, where they produce figs that are eaten by a wide range of bird and mammal species. In our Bornean field site six Ficus seed dispersal guilds can be recognised, differentially attracting subsets of the frugivore community. Guild membership appears to be determined by figs' size, colour, crop size and height above ground, and frugivores' size, sensory and locomotory physiology and foraging height. Vertical stratification therefore appears to be an important determinant of fig and frugivore partitioning. The guild structure observed is discussed with respect to implications for seed dispersal and the differences between the canopy and understorey. Regarding figs eaten primarily by birds, larger fruit and crops can be found in the canopy where they are exposed to larger assemblages of potential frugivores than those presented in the understorey.     

Aspects of variation in a neotropical seed dispersal system

This paper explores the causes and consequences of seed and seedling mortality of the tree Virola nobilis (Myristicaceae) Central Panama in order to understand the advantage to local seed dispersal by birds and monkeys. Post-dispersal mortality due to insects (primarily Conotrachelus spp., Curculionidae) accounts for 30–35% of seed and seedling death during the first 12 weeks after seed fall. Because more seeds and seedlings are killed under and near fruiting trees than 15–45 m away, seed dispersal confers a 20–40 fold advantage on seeds carried 45 m from fruiting adults. In contrast, >60% of seed and seedling death during the first year is due to seed predation by mammals, with >90% due to mammals among the <2% seeds that survive until maternal endosperm is exhausted ±12 weeks after seed fall. Mortaliy due to mammals is independent of distance from parent trees, confering no advantage to seed dispersal. Insects account for variation in mortality attributable to distance effects, mammals to between site effects.Early weevil infestations put a premium on seed removal by large birds (Ramphastos swainsonii, R. sulfuratus, Penelope purpurascens), which carry >50% of the seeds that they eat >40 m, as compared with smaller birds (Baryphthengus martii, Tityra semifasciata, Trogon massena) and monkeys (Ateles geoffroyi), which leave most or all of the seeds that they eat under or near the tree crown.     

Tracking Bat‐Dispersed Seeds Using Fluorescent Pigment1

Tracking the dispersal of seeds by fruit‐eating animals in tropical rain forests is crucial to further our understanding of plant–frugivore interactions and their impacts upon forest regeneration and plant population dynamics. We tested the feasibility of tracking bat‐dispersed seeds in a Philippine lowland rain forest with the help of fluorescent pigment. The powder was mixed with acetone and sprayed to ripe fruits of fig trees, i.e., Ficus septica and F. variegata. During nightly monitoring using a hand‐held ultraviolet lamp bat deposits (seed‐containing spat outs and feces) could successfully be detected. Distances and directions of deposit sites to the focal trees were recorded and seed shadow areas were analyzed. Bats dispersed most of the seeds less than 50 m away from the parent plants resulting in seed shadow areas < 0.30 ha in size. An in situ fruit preference experiment showed that fluorescent powder is unlikely to deter bats from feeding on ripe figs. In conclusion, the technique is simple, inexpensive, noninvasive, applicable to different fields of research and allows one to follow the fate of seeds from known sources.     

Allometric allocation in fruit and seed packaging conditions the conflict among selective pressures on seed size

Selective pressures on seed size could vary among the different stages of plant life cycles, so no simple relation could explain a priori its evolution. Here, we determined the relationships between seed size and two fitness components—seed dispersal and survival from predation—in a bird-dispersed tree, Crataegus monogyna. We interpret these relationships in relation to the patterns of mass allocation to fruit and seed components. Selection patterns were assessed at two levels (1) selection pressures on the parent tree; comparing seed dispersal efficiency among individual plants and (2) selection pressures at the individual seed level; comparing seed size variation (i) before and after dispersal, and (ii) before and after postdispersal seed predation. Dispersal efficiency (percentage of seed crop dispersed) was positively correlated with fruit mass and fruit width. Differences in crop size did not offset this effect, and larger seeds were overrepresented in the seed rain relative to the seed pool before dispersal. However, the advantage of larger seeds during the dispersal stage was cancelled later by an opposite selection pressure exerted by seed predators. As a result, smaller seeds had a higher probability of surviving postdispersal seed predation, establishing an evolutionary conflict imposed by the need for dispersal and the danger of being predated. Birds and rodents preferentially selected highly profitable fruits and seeds in terms of the relative proportion of their components. Larger fruits had a higher pulp to seed proportion than smaller ones, and all seeds had the same proportion of coat relative to the embryo-plus-endosperm fraction. Hence, although predator pressures were stronger than disperser ones, larger seeds invested proportionally less in structural defense than in dispersal.     

Acorn dispersal estimated by radio-tracking

Bird-dispersed seeds are difficult to track, especially in the case of long-distance dispersal events. To estimate the oak dispersal distance and the seed shadow generated by the European jay (Garrulus glandarius), we inserted radio-transmitters in 239 acorns, placed them in bird-feeders and then located them by radio-tracking. Using this methodology we located the exact caching site of 94 Quercus ilex and 54 Q. suber acorns and determined the caching habitat characteristics (vegetation type, distance, spatial distribution). The results show that: (1) there is no differences in the dispersal distance distribution between the different acorn species or sizes, (2) dispersal distances range from approximately 3 m up to approximately 550 m (mean = 68.6 m; median = 49.2 m), (3) recently abandoned fields and forest tracks were the sites preferred by jays to cache acorns, whereas fields and shrublands were avoided and (4) seed shadows showed acorn aggregation zones (i.e. clusters of caches) close to the feeder as well as isolated caches at longer distances. The results also suggest that radio-transmitters are a cheap and reliable way to determine seed shadows and quantify both seed dispersal and post-dispersal seed predation for medium to large seeds.     

Seed rain and advanced regeneration in a tropical rain forest

By comparing seed rain, seedling and sapling community structures we assessed the possible role played by vertebrate seed dispersal as a structuring factor in advanced regeneration of closed-canopied sites in the tropical rain forest of Los Tuxtlas, Mexico. Seed weight, initial morphology of seedlings and species abundance were also analyzed to determine if these traits influenced the probability of establishment in the shade.About half of the seed species falling in five closed forest sites (25×25 m) during one year came from fruiting trees growing within the sites (local seeds) and half from fruiting trees found outside the sites (immigrant seeds). Seeds of liana and upper-canopy species were over-represented among immigrant seeds compared with seeds of understory tree species. This probably reflects the activity of frugivorous arboreal mammals, bats, and birds. Species with immigrant seeds had both a lower abundance and a narrower spatial distribution than locally produced seeds. Therefore, immigrant seeds showed higher diversity values than locally produced seeds.Average seed size and the proportion of epigeous seedlings were similar in local and immigrant species. Under closed-canopied sites, factors affecting community organization seem to operate selectively, favoring the establishment of large-seeded, local abundant species in the advanced regeneration. However, the fact that some saplings of immigrant species were found in the plots suggest that a slow species infiltration may be occurring leading to a slow shift in the advanced regeneration species composition. We propose that the influence of seed dispersal on advanced regeneration structure depends on the disturbance history of the patches where seeds land.     

北京东灵山落叶阔叶林中辽东栎种子雨

在北京东灵山地区的一个落叶阔叶林中调查了辽东栎（Ｑｕｅｒｃｕｓ ｌｉａｏｔｕｎｇｅｎｓｉｓ Ｋｏｉｄｚ．）的种子雨。对于选定的４棵辽东栎中的３棵,树冠下的种子雨分布格局符合二次分布,具有很高的决定系数。由设置在树冠下的种子捕捉器收集的坚果数量来估计整棵树的种子雨。４棵树的种子雨中有活力的种子很少,变化范围从２６到２５９个。每棵树的树冠下的种子雨密度变化范围从０．７６到７．２６个／ｍ＾２。林中平均种     

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).     

Monitoring seed dispersal at isolated standing trees in tropical pastures: consequences for local species availability

The tropical rain forest landscape has been transformed to a mosaic composed of patches of crops, secondary vegetation and remnant forest fragments of different shapes and sizes. Isolation of patches and fragments is a critical issue in the maintenance of local species diversity. In this study we focus on the dispersal of propagules by birds to understant the movement of plants between landscape components. Seed deposition and the behavior of frugivorous birds were monitored at four isolated fig trees (Ficus yoponensis and F. aurea) in man-made pastures. Seed deposition was measured by trapping seeds under canopy trees for six months and by direct observation of bird visits to the four trees for one year. Seed deposition densities were 465, 614, 632 and 1097 seeds/m² accumulated over six months under each of the four trees. We recorded 8268 seeds of 107 species under the trees, among them, 6726 seeds (81%) were of 56 species dispersed by vertebrate frugivores. Seeds of tree species accounted for 26% of the total species. Seventy-three species of birds perched in the observed trees, and 3344 visits were made by 47 species of frugivores. Frugivorous birds occurred in two groups: resident species nesting in the pastures and resident species nesting elsewhere. Propagule exchange between landscape components is clearly influenced by the behavior of these two groups. Structure and dynamics of the landscape depend on plant species availability within the mosaic. This availability is high and suggests possibilities for the management of the local species diversity of tropical rain forests.     

Impact of the spatial uncertainty of seed dispersal on tree colonization dynamics in a temperate forest

An aggregated distribution of dispersed seeds may influence the colonization process in tree communities via inflated spatial uncertainty. To evaluate this possibility, we studied 10 tree species in a temperate forest: one primarily barochorous, six anemochorous and two endozoochorous species. A statistical model was developed by combining an empirical seed dispersal kernel with a gamma distribution of seedfall density, with parameters that vary with distance. In the probability density, the fitted models showed that seeds of Fagaceae (primarily barochorous) and Betulaceae (anemochorous) were disseminated locally (i.e. within 60 m of a mother tree), whereas seeds of Acer (anemochorous) and endozoochorous species were transported farther. Greater fecundity compensated for the lower probability of seed dispersal over long distances for some species. Spatial uncertainty in seedfall density was much greater within 60 m of a mother tree than farther away, irrespective of dispersal mode, suggesting that seed dispersal is particularly aggregated in the vicinity of mother trees. Simulation results suggested that such seed dispersal patterns could lead to sites in the vicinity of a tree being occupied by other species that disperse seeds from far away. We speculate that this process could promote coexistence by making the colonization rates of the species more similar on average and equalizing species fitness in this temperate forest community.     

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.     

海南霸王岭陆均松种子雨和土壤种子库特征

为了解陆均松(Dacrydium pierrei)的自然更新能力,对海南霸王岭陆均松进行为期3年的种子雨、种子库及幼苗幼树观测,并分析其与环境因子的关系。结果表明,陆均松种子雨和种子库有效种子数量太少并存在季节性差异;其结实存在大小年现象;种子雨的扩散密度随着距母树距离的增加而逐渐减少;土壤种子库种子数量在枯枝落叶层所占比例最大,但有活力种子数量极少。幼苗幼树适宜生存的海拔范围为800~1 400 m,经度、纬度、坡度与幼苗幼树分布有明显相关性。因此,选择适宜季节采摘种子,提高种源数量和质量,清除林下枯枝落叶,增加种子萌发能力等以减少陆均松更新的限制条件的人工抚育措施应适当考虑。     

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.     

Aggregated seed dispersal by wreathed hornbills at a roost site in a moist evergreen forest of Thailand

Hornbills (Bucerotidae) are widely regarded as important seed dispersers in tropical forests in Africa and Asia. We investigated how the roosting behavior of wreathed hornbills (Aceros undulatus) influences seed deposition and seedling survival at a roost site in a moist evergreen forest of Khao Yai National Park, Thailand. Fallen fruits and seeds were collected in traps that were placed around a roosting site for 14 months, and seedlings were monitored in adjacent quadrats for 3 years. Seedfall and seedlings of species represented in the hornbill diet occurred at significantly higher densities in the traps and quadrats located beneath the crown of the roosting tree than in those located beyond the crown. With the exception of Cinnamomum subavenium, the seeds and seedlings of most diet species rarely survived beyond the first year. The quality of hornbill dispersal to this roosting site may be poor due to the highly concentrated seedfall, which results in high seed and seedling mortality. However, the number of seeds deposited by each hornbill each day at roosting sites is relatively low. Wreathed hornbills are primarily scatter dispersers during the day and probably serve as agents of seed dispersal in the moist evergreen forest of Khao Yai.     

Seed rain into upland plant communities in Hong Kong, China

Hong Kong is an extreme example of tropical landscape degradation, with no substantial remnants of the original forest cover and a highly impoverished disperser fauna. Seed availability is a potential limiting factor in vegetation recovery in such landscapes. To assess the quantity and quality of the seed rain of woody taxa, seed traps were placed in the major upland vegetation types: fire-maintained grassland, shrubland, and secondary forest. Within the grassland site, traps were placed under isolated trees, isolated male and female shrubs of Eurya chinensis, and in the open. Seeds were collected every 2 weeks for 2 years. The seed rain was highest under female shrubs in grassland (6455 seeds m⁻² year⁻¹), where it was almost entirely confined to their fruiting period. Next highest were isolated trees (890 seeds), followed by male isolated shrubs (611 seeds), shrubland (558 seeds), forest (129 seeds) and open grassland (47 seeds). The number of seed taxa was highest in shrubland (59), followed by isolated trees (42), forest (42), female isolated shrubs (28), male isolated shrubs (15), and open grassland (9). The seed rain differed in species composition between the forest, shrubland, and grassland sites, while the differences within the grassland site were largely in terms of quantity. Birds (particularly bulbuls, Pycnonotus spp.) are known or inferred to be the major dispersal agents for 85% of the seed taxa trapped, 99% of the total number of seeds trapped, and 99.8% of the seeds trapped in the grassland site. Few taxa and of the total seeds were dispersed by wind and no seed taxa were definitely dispersed by fruit bats. The results suggest that even in the most degraded landscape the seed rain is adequate for the development of woody vegetation cover, but that human intervention will be needed for the restoration of plant diversity.     

Seed bank versus seed rain in the regeneration of a tropical pioneer tree

Summary We used the tropical pioneer tree, Cecropia obtusifolia to evaluate the relative importance of different sources of seeds in the regeneration of species that depend on ephemeral sites. We studied seed production in a population established in a 5 ha plot, and dispersal, dormancy and seed predation in two recent treefall gaps (<1 year-old), two building or successional forest patches (10–15 since disturbed), and two mature forest patches (>35 years since disturbed) for a one year period at Los Tuxtlas (Mexico). Flowers and fruits were counted at monthly intervals. Annual fecundity per tree ranged from 1.4×10⁴ to 1.4×10⁷ seeds. Seeds were continuously available on the trees and on the ground. Average annual seed rain per m² (as measured by 0.5×0.5 m seed traps) varied from 184 to 1925 among the six sites. Distance to nearest seed source and patch type explained more than 60% of the seed rain variation among sites. Soil seed density, estimated by counting seeds from ten samples (78.5 cm²×10 cm deep) collected from each site in October and January, ranged among the six sites from 269 to 4485 seeds per m² in January and from 204 to 5073 in October. Soil seed viabilities were much lower (17.1% in October and 5.1% in January) than those of rain seeds (48.26%). Annual survivorships of 2.2% were estimated for seeds artificially sown on the soil surface of a gap and a mature patch, and 3.75% in a building patch. In two other experiments seed removal rates ranged from 27% to 98% in 4 days. Removal rates were significantly higher in gap and mature patches than in building patches. Ants (Paratrechina vividula) and grasshopper nymphs (Hygronemobius. sp.) were the main predators. We draw three main conclusions from our data: (1) Pathogens and predators determine low survivorship of C. obtusifolia's seeds in the soil and a rapid turnover rate (1.07 to 1.02 years) of its seed bank; (2) a continuous and copious seed production and an abundant and extensive seed rain replenish the soil seed pool in patches with different disturbance ages at least up to 86 m from nearest source; (3) more than 90% of the seeds contributing to C. obtusifolia seedling recruitment in gaps are less than one year-old. We discuss our results in the context of previous similar studies for tropical forests.     

Long‐distance dispersal in a fire‐ and livestock‐protected savanna

Savannas are highly diverse and dynamic environments that can shift to forest formations due to protection policies. Long‐distance dispersal may shape the genetic structure of these new closed forest formations. We analyzed eight microsatellite loci using a single‐time approach to understand contemporary pollen and effective seed dispersal of the tropical tree, Copaifera langsdorffii Desf. (Fabaceae), occurring in a Brazilian fire‐ and livestock‐protected savanna. We sampled all adult trees found within a 10.24 ha permanent plot, young trees within a subplot of 1.44 ha and open‐pollinated seeds. We detected a very high level of genetic diversity among the three generations in the studied plot. Parentage analysis revealed high pollen immigration rate (0.64) and a mean contemporary pollen dispersal distance of 74 m. In addition, half‐sib production was 1.8 times higher than full‐sibs in significant higher distances, indicating foraging activity preference for different trees at long distances. There was a significant and negative correlation between diameter at breast height (DBH) of the pollen donor with the number of seeds (r = ?0.640, P‐value = 0.032), suggesting that pollen donor trees with a higher DBH produce less seeds. The mean distance of realized seed dispersal (recruitment kernel) was 135 m due to the large home range dispersers (birds and mammals) in the area. The small magnitude of spatial genetic structure found in young trees may be a consequence of overlapping seed shadows and increased tree density. Our results show the positive side of closed canopy expansion, where animal activities regarding pollination and seed dispersal are extremely high.     

Dispersal of adult female fig wasps 1. Arrivals and departures

Ficus burtt-davyi, like most other fig species (Ficus, Moraceae), is exclusively pollinated by its own unique species of fig wasp, in this caseElisabethiella baijnathi (Chalcidoidea, Agaonidae). Because fig crop development on any one tree is usually synchronised, the small and short-lived female wasps have to migrate and find other trees bearing figs which are at suitable stage of development for oviposition. However, the likelihood of successful location and subsequent arrival at a new host tree is dependent on distance and the effect of environmental factors such as wind and temperature. This study examines the relationship between ambient temperatures and the timing of fig wasps emergence from their natal figs and the commencement of their dispersal flight. The behaviour of the wasps arriving at figs which were ready to be pollinated was also examined. The female wasps did not appear to distinguish between the figs and other parts of the tree when in flight. However, after landing on the tree their search for figs was more directed as they visited more figs than leaves. Short-range recognition of figs appears to be by contact chemo-reception, but the wasps showed a preference for entering figs which did not already contain a female wasp.