Fruit sizes and the structure of frugivorous communities in a New Guinea lowland rainforest

A community of frugivorous insects was studied by rearing of 25 565 individual insects representing three orders (Coleoptera, Lepidoptera and Diptera except Drosophilidae) from 326 woody plant species in a lowland rainforest in Papua New Guinea. Fruits from 19.3% of plant species were not attacked by any insect order, 33.4% of plant species were attacked by a single order, 30% by two orders and 17.2% by all three orders. The likelihood of attack by individual orders was positively correlated so that a higher proportion of plant species than expected suffered either no attack at all or was attacked by all three insect orders. Fruits from most of the plant species exhibited low rates of attack and low densities of insects. One kilogram of fruit was attacked on average by 11 insects, including three to four Coleoptera, six Diptera and one Lepidoptera. Thus, we reared on average one insect from 10 fruits, including one Diptera from 14 fruits, one Coleoptera from 22 fruits and one Lepidoptera from 100 fruits. Only 72 out of the 326 plant species hosted more than one insect per 10 fruits, and only seven species supported a density of greater than one insect per fruit. Our results suggest that specialized insect seed predators are probably too rare to maintain the diversity of vegetation by density‐dependent mortality of seeds as suggested by the Janzen–Connell hypothesis. Fruit weight, fruit volume, mesocarp volume, seed volume and fleshiness had no significant effect on the probability that a fruit would be attacked by an insect frugivore. However, fruits attacked by Diptera were significantly larger and had larger volume of both mesocarp and seeds than fruits attacked by Coleoptera and Lepidoptera.     

Host specificity and interaction networks of insects feeding on seeds and fruits in tropical rainforests

In the tropics, antagonistic seed predation networks may have different properties than mutualistic pollination and seed dispersal networks, but the former have been considerably less studied. We tested whether the structure of antagonistic tripartite networks composed of host plants, insects developing within seeds and fruits, and their insect parasitoids could be predicted from plant phylogenetic distance and plant traits. We considered subsets of the networks (‘subnetworks') at three rainforest locations (Panama, Thailand, Papua New Guinea), based on insect families, plant families or plant functional groups. We recorded 3197 interactions and observed a low percentage of realized interactions, especially in Panama, where insect host specificity was higher than in Thailand or New Guinea. Several factors may explain this, including insect faunal composition, incidence of dry fruits, high fruit production and high occurrence of Fabaceae at the Panamanian site. Host specificity was greater among seed-eaters than pulp-eaters and for insects feeding on dry fruits as opposed to insects feeding on fleshy fruits. Plant species richness within plant families did not influence insect host specificity, but site characteristics may be important in this regard. Most subnetworks were extremely specialized, such as those including Tortricidae and Bruchinae in Panama. Plant phylogenetic distance, plant basal area and plant traits (fruit length, number of seeds per fruit) had important effects on several network statistics in regressions weighted by sampling effort. A path analysis revealed a weak direct influence of plant phylogenetic distance on parasitoid richness, indicating limited support for the ‘nasty host hypothesis'. Our study emphasizes the duality between seed dispersal and seed predation networks in the tropics, as key plant species differ and host specificity tends to be low in the former and higher in the latter. This underlines the need to study both types of networks for sound practices of forest regeneration and conservation.     

Pre-dispersal seed losses to insects in species of Leptospermum (Myrtaceae)

Pre-dispersal seed losses to insects in species of Leptospermum at Wilson's Promontory in south-eastern Australia varied markedly between species, seasons, sites and individuals. The proportion of fruits attacked by insects averaged 11% (range 3–37%) in populations of Leptospermum juniperinum, 23% (6–95%) in Leptospermum myrsinoides, 23% (0–47%) in Leptospermum laevigatum, and 28% (8–45%) in Leptospermum lanigerum. The proportion of seeds lost in attacked fruits also varied widely. Differences between species varied markedly from year to year, but differences between sites were relatively constant. Fruit production by L. myrsinoides was unusually low at most sites during 1984; in each case this coincided with unusually high rates of insect attack, suggesting that predator satiation operates during normal seasons.     

Interaction between juniper Juniperus communis L. and its fruit pest insects: Pest abundance, fruit characteristics and seed viability

The relationships between the fruit features of Juniperus communis and the presence of fruit pests were studied in Sierra Nevada, SE Spain. The abundance of two insect species — a pulp-sucking scale and a seed-predator wasp — was surveyed with respect both to fruit characteristics and to viability of seeds contained therein. Seed-predator pressure was not significantly related to any fruit characteristics; however, pulp suckers tended to be more abundant in plants with low pulp: seed ratios and high fruit-water content. In addition, fruits with high levels of pulp-sucker attack tended to have higher water content. A multi-factor ANOVA, considering the identity of the plant and the attack of the different pests as factors, showed that plant identity accounts for most of the variation in fruit characteristics. The viability of seeds tended to be lower in plants strongly attacked by both pests. Fruits attacked by seed predators showed significantly lower proportions of viable and unviable seeds than did unattacked fruits. Seed viability was also lower in those fruits heavily attacked by pulp suckers, but this pattern is strongly mediated by plant identity. Pest activity proved to be clearly associated with a direct decrease in juniper reproductive capacity. This loss involved a reduction of the viable-seed number, mainly related to the seed predator, as well as a reduction of fruit attractiveness to frugivorous dispersers, related to the pulp sucker.     

A highly resolved food web for insect seed predators in a species‐rich tropical forest

The top‐down and indirect effects of insects on plant communities depend on patterns of host use, which are often poorly documented, particularly in species‐rich tropical forests. At Barro Colorado Island, Panama, we compiled the first food web quantifying trophic interactions between the majority of co‐occurring woody plant species and their internally feeding insect seed predators. Our study is based on more than 200 000 fruits representing 478 plant species, associated with 369 insect species. Insect host‐specificity was remarkably high: only 20% of seed predator species were associated with more than one plant species, while each tree species experienced seed predation from a median of two insect species. Phylogeny, but not plant traits, explained patterns of seed predator attack. These data suggest that seed predators are unlikely to mediate indirect interactions such as apparent competition between plant species, but are consistent with their proposed contribution to maintaining plant diversity via the Janzen–Connell mechanism.     

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.     

Seed characteristics and susceptibility to pathogen attack in tree seeds of the Peruvian Amazon

Many studies now suggest that pathogens can cause high levels of mortality in seeds and seedlings. Recruitment from seed to sapling is an important bottleneck for many tree species, and if specialist or generalist pathogens have differential negative effects among species of juvenile trees, then they may have a significant impact on forest community structure. To explore the effects of differential pathogen attack among tropical tree species, we quantified pathogen attack on the seeds of 16 tree species from the southeastern Peruvian Amazon and asked which seed characteristics, including size, hardness, germination time and mode, shade tolerance, and fruit type, were most closely correlated with susceptibility to pathogens. Shade tolerance and seed weight were positively and significantly correlated with susceptibility to pathogen attack by ecological trait regressions (ETRs), and correspondence analysis indicated that there might be increased attack rates in species with brightly colored, pulpy fruits (often dispersed by primates). Only shade tolerance was significantly correlated with pathogen attack when the analyses accounted for phylogenetic relatedness between species. Thus, contrary to standard predictions of size-defense ratios, our results suggest that larger, shade-tolerant seeds tend to be more susceptible to pathogen attack than smaller, light-dependent seeds. Moreover, differential pathogen attack may shape seed community composition, which may affect the successful recruitment of adults.     

Predispersal seed-predation by insects in the Venezuelan Central Plain: Overall patterns and traits that influence its biology and taxonomic groups

Predispersal seed predation (PSP) by insects was studied in a plant community of the Venezuelan Central Plain (VCP). The main goal was to examine to what extent vegetation structure and fruit–seed attributes determined the incidence of PSP by insects at the community level. We studied a total of 187 species from 59 families, in five habitat types. The proportion of seed-predated plant species (N=89; 47.6% of the total) was explained by different factors such as the abundance of legume families, plant species richness, fruit dehiscence, seed biomass, and starch content. Coleoptera was the most diverse taxonomic order of insect seed predators, with Bruchidae and Curculionidae showing the largest number of genera and species, followed by Lepidoptera (Pyralidae). Bruchidae and Pyralidae, but not Curculionidae, tended to be separated according to vegetation attributes, such as vegetation structure and flowering and fruiting phenology. In addition, Bruchidae was associated with dehiscent fruits, legumes, epizoochory, and granivory, whereas Curculionidae was more related with indehiscent fruits and endozoochory, and Pyralidae with abiotic seed dispersal. Bruchid larvae tended to prey upon single seeds, whereas larvae of Lycaenidae usually preyed upon more than one seed per fruit; Curculionidae did not show any clear pattern on this. One larva developing in a single seed (Type I) was the most common type of PSP, closely followed by one or more larvae developing outside seeds within the fruit (Type II). Type III (seed predation by adult insects inside the fruit) was the least common. Type I appears to occur most frequently in climbers, in the forest–savanna transition habitat, and in those cases in which insect adults emerged during the rainy season. By contrast, Types II and III tended to be associated with annual herbs, the lowest stratum (0.05–0.6 m), and disturbed areas. Moreover, Type I was usually found in samaras, drupes, and indehiscent fruits, whereas Type II and Type III were more commonly observed in capsules. Type I and Type II seeds differed in their nutritional composition, which in turn appears to affect seed predation specialization. The number of insect seed predator species and plant richness per habitat were positively correlated. The number of seed predator species was associated with the abundance of trees and climbers because more complex plant life forms offer a large variety of resources, woody species frequently had fruits and seeds larger than herbaceous species, and trees and climbers showed the lowest values of host specificity. To our knowledge, this study is the first one that characterizes the community of predispersal seed predators and simultaneously evaluates ecological, morphological, and nutritional factors determining the groups of seed predators and how PSP occurred.     

Is the inverse leafing phenology of the dry forest understory shrub Jacquinia nervosa (Theophrastaceae) a strategy to escape herbivory?

In the dry forest of Santa Rosa National Park, Costa Rica, the understory shrub Jacquinia nervosa presents an inverse pattern of phenology that concentrates vegetative growth and reproduction during the dry season. In this study, we tested the "escape from herbivory" hypothesis as a potential explanation for the inverse phenological pattern of J. nervosa. We monitored leaf, flower and fruit production in 36 adult plants from October 2000 to August 2001. Leaves of six randomly selected branches per plant were marked and monitored every two weeks to measure the cumulative loss in leaf area. To analyze pre-dispersal seed predation we collected 15 fruits per plant and counted the total number of healthy and damaged seeds, as well as the number and type of seed predators found within the fruits. Leaf, flower, and fruit production occurred during the first part of the dry season (end of November to February). The cumulative herbivory levels were similar to those observed in other tropical dry forest tree species that concentrate leaf production during the wet season, and were concentrated on young leaves, which lost an average of 36.77 % of their area (SD = 34.35 %, N = 195). Chewing beetles of the genus Epicauta (Meloidae) were the most important herbivores. In mature leaves, most of the damage was caused by the beetle Coptocycla rufonotata (Chrysomelidae). Fruits took 4 months to develop during the dry season (January-March 2001) but continue increasing in size well into the first 3 months of the wet season (May-July). Average seed number per ripe fruit was 9 (SD = 5, N = 500). Seed predation in mature fruits was 42 % (SD = 47 %, N = 122). Most seeds were damaged by moth larvae of the family Tortricidae. Only 3 % of the flowers became fruits. This was influenced by the low level of flower synchrony (0.38+/-0.26, N = 36 plants), but neither leaf synchrony (0.88+/-0.06, N = 36 plants) nor plant size influenced fruit numbers. The significant damaged produced by insect herbivores in young leaves, fruits, and seeds, as well as the low reproductive index observed in J. nervosa, shows that the inverse leafing phenology of this species is not consistent with the "escape hypothesis" since J. nervosa was considerably attacked during the dry season. Considering the strong seasonality of the tropical dry forest and the heliophyte character of J. nervosa, it is more likely that this phenological strategy evolved in response to seasonal fluctuations in light availability, light quality, and daylength.     

Seed predation by insects in tropical mangrove forests: extent and effects on seed viability and the growth of seedlings

Summary Although insects are known to be important seed predators in most terrestrial forests, their role in marine tidal (mangrove) forests has not been examined. Surveys at 12 sites in tropical Australia showed that between 3.1 and 92.7 percent of the seeds or propagules of 12 mangrove tree species had been attacked by insects. Seeds/propagules of six species (Avicennia marina, Bruguiera gymnorrhiza, B. parviflora, Heritiera littoralis, Xylocarpus australasicus and X. granatum) showed consistently high (>40%) levels of insect damage. Greater than 99% of H. littoralis seeds were attacked by insect predators. The survival and subsequent growth in height and biomass of insect-damaged and non-damaged control seeds/propagules of eight mangrove species were compared in shadehouse experiments. Mangrove species fell into 4 groups with regard to the effect of insect predators on their seeds and seedlings. Xylocarpus australasicus and X. granatum had significantly decreased survival (X 48 and 70%) and growth in height (X 61 and 96%) and biomass (X 66 and 85%). Bruguiera parviflora showed decreased survival (X 59%), but there was no effect of insects on the growth of surviving propagules. In contrast, there was no effect of insect damage on the survival of seedlings of Avicennia marina and Bruguiera exaristata, but decreased growth in height (X 22 and 25%) and biomass (X 22 and 26%). Survival and growth of seedlings of Rhizophora stylosa and Bruguiera gymnorrhiza were not affected. The influence of insect seed predators on the survival and growth of seeds of mangrove species in forests will depend on the relative abundance of seed-eating crabs and intertidal position in mangrove forests.This is Contribution No 499 from the Australian Institute of Marine Science     

Performance of the fir coneworm Dioryctria abietivorella (Grote) as affected by host species and presence or absence of seed cones

1 Larval performance of Dioryctria abietivorella (Grote) (Lepidoptera: Pyralidae) was compared when insects were reared in white spruce, black spruce and Jack pine seed orchards. For each species, half of the insects developed in the presence of cones, while the other half was maintained on branches without cones. 2 Significantly faster development rates were observed on spruce species when compared to Jack pine. Significant differences in survival were also recorded between the different feeding treatments. 3 The presence of seed cones on spruce species significantly increased insect mean weight compared with branches without cones, but no significant differences were noticed between the different tree species with cones. 4 White spruce is the most suitable host tree for fir coneworm feeding and the availability of seed cones plays an important role in determining D. abietivorella larval performance.     

Occurrence of Amblycerus species in Cordia trichotoma seeds and their influence on germination

Forest species can have their seeds damaged by granivorous insects, especially by those in their larval stage. In this context, this study aims to report the occurrence of Amblycerus species in Cordia trichotoma seeds, to describe their main damage to seeds and effects on germination, as well as their associated hymenopteran parasitoids. Therefore, seven trees were selected in the municipality of Taquaruçu do Sul, RS, Brazil. Fruits were collected weekly from the medium third of the tree crown, from the beginning of their formation until total dehiscence. To examine the damage caused by granivorous insects within the fruits, 15 fruits from each tree were sectioned with a scalpel. Furthermore, 10 fruits from each tree were stored individually in clear plates to verify the occurrence and identification of granivorous insect species. Evidence of the damage caused to seeds was verified through the germination test by comparing preserved and damaged seeds, with four repetitions of 25 seeds each. The species Amblycerus longesuturalis and Amblycerus profaupar (Chrysomelidae: Bruchinae) were found associated with fruits of C. trichotoma. Female insects predominantly laid eggs on the superior part between the marcescent calyx and the fruit, and larvae perforated the fruit tegument to start consuming seed embryos and reserves. Bruquine larvae are parasitized by Hymenoptera of Bracon, Mirax, Omeganastatus and Triapsis genera. In conclusion, the germination of C. trichotoma seeds is significantly affected by emergence orifices caused by granivorous species.     

Phytophagous insect-woody sprout interactions in tropical eucalypt forest. II. Insect community structure

Abstract Sucking insects constituted 79% of all phytophagous insects collected from woody sprouts in the ground layer of a tropical eucalypt forest. Mobile insect groups such as non-psyllid Hemiptera and Orthoptera were relatively frequent in this environment compared to temperate, Eucalyptus-dominated vegetation. The high fire frequency of the tropical eucalypt forest may favour mobile insect groups. The capture of sucking insects and caterpillars peaked in dry season samples. Other patterns of abundance of phytophagous insect groups showed little consistency in their seasonal trends between host species or between vegetation types within host species. Disparities between chewing insect abundance in daytime samples and the damage chewing insects cause, may result from disproportionate consumption by large, mainly nocturnal insects, such as members of the Orthoptera. In this study, 21% of insect species were specialists on single plant species. This study suggested that insect abundance reflected the growth patterns of woody sprouts after regular burning, rather than that plant growth and development were tuned to the pressures of insect herbivory.     

Differences in the fruit maturation stages at which oviposition occurs among insect seed predators feeding on the fruits of five dipterocarp tree species

The seeds of dipterocarp trees are the main food resources for many species of weevils, bark beetles and small moths; however, for most seed‐eating insects on dipterocarp tropical trees, seed utilization patterns remain poorly investigated. This study aimed to determine the fruit maturation stages at which eggs are laid by different insect seed predators feeding on the seeds or fruits of the following five dipterocarp species: Dipterocarpus globosus, Dryobalanops aromatica, Shorea beccariana, S. acuta and S. curtisii, which reproduced during the same period. We investigated the occurrence frequencies of the insect seed predators at various growth stages by collecting both unfallen and fallen fruit on several occasions during the period of seed/fruit maturation in a tropical rainforest in Borneo from September to December 2013. Weevils and bark beetles were the dominant insect seed predators of the five tree species. One or two weevil species of Alcidodes, Damnux and/or Nanophyes preyed on the seeds of each of the five tree species, and one bark beetle species, Coccotrypes gedeanus, preyed on the seeds of all five tree species. Many larvae, pupae and adults of each weevil species were found in pre‐dispersal (unfallen) fruit, whereas bark beetles at various growth stages were found in post‐dispersal (fallen) fruit. These results suggested that, among the dominant insect seed predators of the five dipterocarp species, weevil species oviposit on pre‐dispersal fruit and begin their larval growth before seed dispersal, whereas the oviposition and larval development of bark beetle species occurs in post‐dispersal fruit.     

Can remnant frugivore species effectively disperse tree seeds in secondary tropical rain forests?

Seed dispersal by frugivores in tropical rain forests is important for maintaining viable tree populations. Over the years, vertebrate assemblages in tropical forests have been altered by anthropogenic disturbances, leading to concerns about the ability of remnant vertebrates to substitute for the lost or declining vertebrate populations. We compared vertebrate composition and frugivore visitation rates as an indirect measure of rate of seed dispersal in three tropical rain forests in Uganda, namely Mabira, Budongo and Kibale Forests. Mabira is highly disturbed, Kibale is little and Budongo is intermediate. The aim was to determine whether vertebrate assemblages in differentially disturbed forests had comparable abilities to disperse seeds and whether tree species were equally vulnerable to loss of seed dispersers. Assemblages of forest generalist species were similar in all forests, but specialists were less abundant in the heavily disturbed forest. Remnant frugivores in the heavily disturbed forest were mainly small-bodied species that spat seeds beneath fruiting trees compared to large-bodied species observed in the less disturbed forests that ingested and carried away the seeds. We postulate that the quantity of seeds dispersed in heavily disturbed forests is much reduced due to low visitation rates of frugivores and the absence of large frugivores that consume large quantities of fruit. The quality of seed dispersal is affected as well by the distance over which seeds are moved. Assessment of vulnerability of trees shows no evidence for disperser substitution for trees producing large fruits. Fruit trees with low nutritional contents and digestibility were least visited in frugivore-impoverished forests. The loss of large specialist frugivores is likely to affect recruitment of many trees, especially of species that cannot establish beneath adult conspecifics.     

The insect‐focused classification of fruit syndromes in tropical rain forests: An inter‐continental comparison

We propose a new classification of rain forest plants into eight fruit syndromes, based on fruit morphology and other traits relevant to fruit‐feeding insects. This classification is compared with other systems based on plant morphology or traits relevant to vertebrate fruit dispersers. Our syndromes are based on fruits sampled from 1,192 plant species at three Forest Global Earth Observatory plots: Barro Colorado Island (Panama), Khao Chong (Thailand), and Wanang (Papua New Guinea). The three plots differed widely in fruit syndrome composition. Plant species with fleshy, indehiscent fruits containing multiple seeds were important at all three sites. However, in Panama, a high proportion of species had dry fruits, while in New Guinea and Thailand, species with fleshy drupes and thin mesocarps were dominant. Species with dry, winged seeds that do not develop as capsules were important in Thailand, reflecting the local importance of Dipterocarpaceae. These differences can also determine differences among frugivorous insect communities. Fruit syndromes and colors were phylogenetically flexible traits at the scale studied, as only three of the eight seed syndromes, and one of the 10 colors, showed significant phylogenetic clustering at either genus or family levels. Plant phylogeny was, however, the most important factor explaining differences in overall fruit syndrome composition among individual plant families or genera across the three study sites. Abstract in Melanesian is available with online material.     

Seed Dispersal and Ingestion of Insect-Infested Seeds by Black Howler Monkeys in Flooded Forests of the Parana River, Argentina

All howler monkey species ( Alouatta spp.) have a folivorous–frugivorous diet. Howler monkeys are reported to be seed dispersers in several areas, including black howlers ( Alouatta caraya ), which are important seed dispersers in northern Argentinean forests. The goal of this work was to study the three-way interaction between insects, seeds, and black howlers, and assess the functional significance of this tri-trophic interaction for seed dispersal. I determined through direct observation that fruits of species with a high proportion of insect infestation were important components of howler monkey diet. Ocotea diospyrifolia seeds from fresh faeces of black howlers contained dead larvae, but seeds were still able to germinate. Seeds in which larvae had reached an advanced stage of development did not germinate. Larvae of infested Eugenia punicifolia fruits were killed by digestion when they occurred in the pulp early in the fruiting season, but were dispersed alive with seeds later in the season. Banara arguta fruits contained both healthy and infested seeds; infested seeds were destroyed during digestion, while healthy seeds were dispersed. Black howlers' ingestion of infested fruits could result in the: (1) killing of larvae and dispersion of healthy seeds; (2) spread of larvae; or (3) destruction of infested seeds. This will depend on the relationship between the time at which fruit is consumed by black howlers, the time at which insect infestation occurs, and also probably on the hardness of the seed coat and the seed–insect size ratio.     

Empty seeds reduce seed predation by birds in Juniperus osteosperma

Utah juniper (Juniperus osteosperma) is one of many plant species that produce large numbers of fruits containing parthenocarpic or otherwise empty or inviable seeds. We tested the hypothesis that production of empty fruits in this species results in reduced levels of predation on fertile seeds. In a population in west-central Utah, we estimated the proportion of fruits with filled seeds in trees suffering high levels of fruit destruction by the seed-eating bird Parus inornatus and in neighbouring trees similar in crown and fruit-crop size but suffering negligible predation. We found that the heavily attacked trees had higher proportions of filled seeds. Thus, juniper may benefit from producing fruits that contain no offspring. This is the first study to demonstrate that empty seeds may reduce predation by vertebrate seed eaters and the first to demonstrate discrimination based on seed filling at the level of whole plants.     

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.     

Soil seed banks in dry Afromontane forests of Ethiopia

Abstract. The soil seed bank was investigated in four dry Afromontane forests of Ethiopia. At least 167 plant species were identified in the 0–9 cm soil layer with total densities ranging between 12 300 and 24 000 seeds/m². Herbs were represented with the largest numbers of species and seeds in the seed bank, while the contribution of tree species was generally low. The overall vertical distribution of seeds was similar at all sites with the highest densities occurring in the upper three cm of soil and gradually decreasing densities with increasing depth. Relatively high densities also occurred in the litter layer. There were large differences in depth distribution between species, suggesting differences in seed longevity. A large number of species in dry Afromontane forests evidently store quantities of seeds in the soil and this is in contrast to the situation in most tropical rain forests, dry lowland forests and savannas, where both the number of seeds and the number of species are relatively small. It is possible that the strongly seasonal and unpredictable climate of this region may have selected for high levels of dormancy, and that herb regeneration is associated with small scale disturbance. The fact that most of the dominant tree species do not accumulate seeds in the soil suggests that their regeneration from seed would be unlikely if mature individuals disappeared. Most tree species have relatively large seeds and poor long-distance dispersal; this implies that restoration of Afromontane forests after destruction would be difficult. Since there is a diverse seed bank of the ground flora, this component of the vegetation would have a better chance of reestablishing. However, because most cleared forest land is used for agricultural crop production, it is probable that the seed bank will be depleted in only a few years. Therefore, the future of the Afromontane forest flora seems to depend on the successful conservation of the few fragments of remaining natural forest.