Predispersal seed predation on five Piper species in tropical rainforest

Absolute number of seeds lost to predispersal seed predators and proportion of total seeds lost per infructescence were compared among five Costa Rican Piper species of different annual fecundities. Mean seed number and mean seed size in the five species were negatively correlated. The impact of predation on these species was inversely related to the number of seeds they produced. The two early successional species had very high fecundities, a combination of many seeds per infructescence, many infructescences per plant, and, in one species, year-round reproduction. Although seed predators destroyed as many or more seeds of these early successional species than they did of the less fecund, late successional species, this loss accounted for a relatively minor proportion (9 and 12%) of the seeds of the early successional species. In contrast, late successional species produced fewer, larger seeds in a smaller number of infructescences and were not continually in fruit. One of these species, which produced intermediate numbers of intermediately sized seeds, lost 30% of the seeds in each infructescence on average. Seed predators destroyed a larger proportion (65 and 76%) of the seeds per infructescence in the two species with fewest seeds per infructescence. High levels of insect damage in these late successional species caused many of their infructescences to abort prematurely. Taken together these factors resulted in annual fecundities several orders of magnitude smaller in shade-tolerant Piper species than the annual fecundities of shade-intolerant, early successional species. Seedlings of the two early successional species were common in large gaps and other sunny clearings and seedlings of the species with 30% seed loss were occasional, whereas no seedlings were seen of the two species with the highest proportional seed loss, suggesting that seed predation on the latter species may limit seedling recruitment.     

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.     

Biogeography of seed-dispersal syndromes, life-forms and seed sizes among woody rain-forest plants in Australia's subtropics

Aim To enhance our understanding of the evolutionary interactions between seed‐dispersal syndromes, life‐forms, seed size, and habitat characteristics by studying their association with the regional‐scale distributions of subtropical rain‐forest plants in the context of climatic gradients. Location South‐east Queensland, subtropical eastern Australia (152° E, 26° S). Methods We classified 250 rain‐forest sites into six floristic site‐groups based on their woody plant composition. The resulting classification was strongly associated with variation in rainfall. The distribution of species across the floristic site‐groups was used to assign 568 species to seven habitat classes (one class for ‘widespread’ species, with all other species classified according to the site‐group within which they were most frequent). Species were also classified for three other categorical life‐history factors: three dispersal syndromes based on diaspore morphology (fleshy, wind‐assisted, and unadorned); four life‐forms (trees, shrubs and small trees, tall climbers, and short and shrubby climbers); and four seed‐diameter classes (< 3 mm, ≥ 3 and < 4.5 mm, ≥ 4.5 and < 7 mm, and ≥ 7 mm). We used a basic comparative approach augmented by simple phylogenetically constrained comparisons to assess association between dispersal syndrome, seed size, life‐form, and habitat class. Results Across the rain forests of south‐east Queensland, the proportion of species with fleshy diaspores or of large stature increases with rainfall. High‐rainfall sites also have larger average seed sizes, but the difference in average seed size between high‐ and low‐rainfall sites is small compared with variation within sites. Among species, those with fleshy fruit tend to have larger seeds and to favour high‐rainfall sites. Very few small trees produce diaspores adapted for wind‐assisted dispersal. On average, species with unadorned diaspores have smaller seeds than those with fleshy diaspores. However, within sites, species with unadorned and fleshy diaspores have similar average seed sizes, and some species with unadorned diaspores from high‐rainfall habitats have extremely large seeds. Main conclusions Commonly observed associations between fleshy fruit, larger plants, larger seeds, and productive habitats are apparent within the rain‐forest flora of south‐east Queensland. However, these associations are generally weak and involve complex interactions. For example, the strong tendency for species with fleshy fruit to have larger seeds than those with unadorned diaspores concealed a significant group of species from wetter forests that produce extremely large seeds and unadorned diaspores. The most widespread species in this study tend to be large plants (particularly robust lianes) and to produce fleshy fruit, but they tend not to have relatively large seeds. The association between large seeds, large plants, fleshy fruit and productive habitats is discussed as part of an evolutionary strategy favouring fitness in populations close to carrying capacity. We review some problems with focusing on establishment chances per seed as the driver towards association between large seeds, large plants and productive rain‐forest habitats (the difficult‐establishment hypothesis). Instead we suggest that production of large, short‐lived seeds by long‐lived plants in temporally stable, closed habitats may reflect the limited evolutionary potential for strategies enhancing colonization (e.g. producing large numbers of dormant seeds), thus allowing the establishment benefits of large seeds greater selective influence (the slow‐replacement hypothesis). The association of fleshy fruit with large seeds probably reflects the difficulty of dispersing large seeds by other means (the difficult‐dispersal hypothesis).     

Small mammals as potential seed dispersers in New Zealand

Weed invasion success is strongly influenced by availability of seed dispersal vectors, which may include animals. We examined the potential of several small introduced mammals (mice, kiore, ship rats and possums) to disperse germinable seeds in New Zealand. Captive animals were fed fleshy fruit of weeds (Berberis glaucocarpa, Cotoneaster spp., Crataegus monogyna, Ilex aquifolium, Leycesteria formosa, Ligustrum sinense, Lonicera japonica, Passiflora mollissima, Pyracantha angustifolia, Sorbus hupehensis) and native species (Coprosma spp., Prumnopitys ferruginea and Solanum aviculare). We recorded the percentage of fruit consumed, seed ingested and gut passage time. Faeces were collected and the seeds extracted and tested for germination potential in an unheated glasshouse (two weed species) or under controlled conditions (11 species). The smallest rodents (mice and kiore) generally destroyed all seeds eaten. Large numbers of viable seeds of the small‐seeded (<1 mg) species, L. formosa and S. aviculare, passed through ship rats. Possums consumed the seeds of all adventive and native fruits except P. ferruginea. The proportion of seeds recovered intact from possum faeces varied with plant species and ranged from 6 to 83%. The time required for 50% of all seeds to be passed by possums ranged from 2.5 to 5.5 days with an average of 3.7 days, and was generally unrelated to simple fruit parameters such as percentage pulp and moisture content. For seeds where germination also occurred in the uneaten controls, the germination of seed from possums ranged from 3 to 78%. Germination was mostly lower in seeds from possums than in the controls, where differences were significant. Possums have major potential to disperse a wide range of fleshy fruit‐producing native and introduced plant species. Ship rats have the potential to disperse those with very small seeds.     

Barriers to Forest Regeneration in an Abandoned Pasture in Puerto Rico

Sources of forest regeneration (soil seed bank, seed rain) and barriers to seedling establishment were examined in a recently abandoned pasture in eastern Puerto Rico. Few woody species were found in the soil seed bank or in the seed rain. The number of seeds and species in the seed rain and soil seed bank declined with distance from the adjacent secondary forest. Nine species naturally dispersed and colonized plots during the study, with the wind‐dispersed tree Tabebuia heterophylla being the predominant colonizer (91% of all seedlings). Barriers to seedling establishment were determined using a blocked field experiment with eleven woody species representative of a variety of life histories. Each species was planted under the pasture vegetation (control) or in areas where all vegetation was removed (removal). Germination was enhanced for four species in the control treatment, five species were not affected, and two species did not germinate under either treatment. Survival to 6 months was higher in the removal treatment for two species. Seedling biomass was greater in the removal treatment at 12 months for one species. Seed mass was a good predictor of germination success and final shoot biomass, but not survival. This study demonstrates that seeding recently abandoned pastures with a mix of known pioneer species may accelerate the rate of secondary succession, but some species will have to be planted in later successional stages in order to overcome strong barriers to establishment.     

Role of small rodents in the seed dispersal process: Microcavia australis consuming Prosopis flexuosa fruits

Understanding the functional role of animal species in seed dispersal is central to determining how biotic interactions could be affected by anthropogenic drivers. In the Monte Desert, mammals play different functional roles in Prosopis flexuosa seed dispersal, acting as opportunistic frugivores (endozoochorous medium‐sized and large mammals) or seed hoarders (some small sigmodontine rodents). Our objective was assessing the functional role of Microcavia australis, a small hystricognathi rodent, in the fruit removal and seed deposition stages of P. flexuosa seed dispersal, compared to sympatric sigmodontine rodents. In situ, we quantified fruit removal by small rodents during non‐fruiting and fruiting periods, and determined the distance seeds were transported, particularly by M. australis. In laboratory experiments, we analysed how M. australis stores seeds (through scatter‐ or larder‐hoarding) and how many seeds are left in caches as living seeds, relative to previous data on sigmodontine rodents. To conduct field studies, we established sampling stations under randomly chosen P. flexuosa trees at the Ñacuñán Man and Biosphere Reserve. We analysed fruit removal by small rodents and seed dispersal distance by M. australis using camera traps focused on P. flexuosa fruits covered with wire screen, which only allowed entry of small animals. In laboratory trials, we provided animals with a known number of fruits and assessed seed conditions after removal. Small rodents removed 75.7% of fruit supplied during the non‐fruiting period and 53.2% during the fruiting period. Microcavia australis and Graomys griseoflavus were the main fruit removers. Microcavia australis transported seeds to a mean distance of 462 cm and cached seeds mainly in scatter‐hoards, similarly as Eligmodontia typus. All transported seeds were left in fruit segments or covered only by the endocarp, never as predated seeds. Microcavia australis disperses P. flexuosa seeds by carrying fruits away from a source to consume them and then by scatter‐hoarding fruits and seeds.     

Importance of Seed and Microsite Limitation: Native Wildflower Establishment in Non‐native Pasture

Barriers to establishing native plant communities on former pasture include dominance by a single planted species, hydrologic and edaphic alteration, and native species propagule limitation. Establishment may be dispersal‐limited (propagules do not arrive at the site), microsite‐limited (areas suitable for seedling emergence and survival do not exist), or both. Successful restoration strategies hinge on identifying and addressing critical limitations. We examined seed and microsite limitation to establishment of a native wildflower (Coreopsis lanceolata ) in a former pasture dominated by Paspalum notatum (bahiagrass). We determined the relative and interactive effects of microsite (irrigation and disturbance) and seed limitation on C. lanceolata establishment. We tested (1) irrigation (none, pre‐seeding, and pre‐ and post‐seeding), (2) disturbance (none, sethoxydim, glyphosate, and topsoil removal), and (3) C. lanceolata seeding rate (three seeding densities). Applying glyphosate before seeding increased C. lanceolata establishment compared to other disturbance treatments. Ultimately, C. lanceolata establishment was not affected by irrigation. Coreopsis lanceolata establishment was limited when seeded at 100 live seeds/m² but not at 600 or 1100 live seeds/m². Seed and microsite availability interactively affected C. lanceolata establishment, in that microsite limitation was biologically relevant only when a minimum number of seeds were present. In practice, both seed and microsite requirements must be met for successful establishment, and increasing the availability of seeds or microsites does not compensate for limitations of the other. Here, it is the relative importance of seed and microsite limitations that drives plant establishment; these limitations do not represent a simple dichotomy.     

Comparing methods of weed seed exposure to predators

Several methods of seed exposure are used in seed predation studies, but how these methods influence the results remains poorly studied. In this article, two commonly used methods of seed exposure – seed cards and plasticine trays – were compared in the field and in the laboratory using three species of weed seeds. In the field, cards or trays with seeds were exposed either with or without cages to keep vertebrates out and either with or without impermeable roofs to provide protection from the weather. The overall seed retrieval from the control stands varied significantly between the methods of seed exposure, roof treatment and the species of seeds, and the scatter in the seed retrieval increased or decreased monotonically with the temperature or precipitation for some species of seeds. This indicates that the controls were more or less relevant depending on the weather conditions and species. The seed removal from cards varied between the species of seeds. The effect of exclosure cages indicated that invertebrates were the dominant seed predators of Capsella bursa‐pastoris, while in Poa annua and Stellaria media both vertebrates and invertebrates were important. Higher seed removal from plasticine trays compared to seed cards was found for all three species of seeds, and placing roofs over the seeds affected seed removal in C. bursa‐pastoris and P. annua. In the laboratory, seed consumption varied with the method of exposure in 10 out of 12 combinations of seed species and predator (two species of carabids and two species of isopods). The overall tendency was the opposite of the field observations: seed consumption was higher for seeds on filter paper and cards compared to seeds on tin trays. We conclude that seed cards are more convenient for short term studies in the field, while tin trays are useful in laboratory multi‐choice experiments. To measure the realistic consumption of invertebrate predators in the laboratory, filter paper seems to be the best option as it does not present an obstacle to predators eating the seed. Using roofs introduced further bias regarding the estimates of seed removal in the field and thus should be avoided.     

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.     

Seed Removal Rates Under Isolated Trees and Continuous Vegetation in Semiarid Thornscrub

Clearing native vegetation to increase the amount of land available for agriculture in northeastern Mexico has left remnants ranging in size from fragments of continuous vegetation to isolated individual trees. These provide valuable opportunities for restoring larger areas of native vegetation. We explored whether fragmentation of Tamaulipan thornscrub affects the removal of seeds from 12 woody species that encompass a range of sizes and dispersal mechanisms. We tested whether (1) seed removal rates under isolated trees were higher than under continuous vegetation; (2) dispersal structures, such as fleshy pericarps, made some seeds more attractive to seed removers; and (3) microenvironmental variation affected seed removal rates. Seeds were placed under canopies of Texas ebony (Ebenopsis ebano) and Mesquite trees (Prosopis laevigata). Seed removal trials were conducted three times, each trial lasting 30 days. Most seeds were removed in all trials by the end of one month. Seed removal rate was slower under isolated trees. In general, fleshy fruits were removed faster than other fruits; whole fruits and fleshy tissue were removed faster than depulped seeds. In species with fleshy pericarps, acid washing of seeds, to simulate seeds processed in the digestive tract of dispersers, reduced the seed removal rates, suggesting that it would be a good pre‐treatment for restoration efforts.     

The role of frugivorous birds in fruit removal and seed germination of the invasive alien Cotoneaster franchetii in central Argentina

Many invasive plant species have fleshy fruits that are eaten by native frugivorous birds which disperse their seeds and may facilitate their germination, playing an important role in plant invasion success. The fleshy‐fruited shrub Cotoneaster franchetii (Rosaceae) is an important invasive alien in the mountainous regions of central Argentina. To determine the role of avian frugivorous in fruit removal of this species, we conducted a frugivore exclusion experiment including bagged and unbagged branches in 75 plants of C. franchetii. At the end of the dispersal period, we compared the percentage of missing fruits (removed by birds + naturally dropped) in unbagged branches with the percentage of naturally dropped fruits in bagged branches. To assess whether any mechanism acting on seeds during their passage through bird guts (de‐inhibition by pulp removal and/or seed scarification) affects seed germination of this species, we compared percentage and speed of germination among seeds obtained from faeces of the native frugivorous Turdus chiguanco, from manually de‐pulped fruits, and from intact fruits. The percentage of missing fruits per shrub in unbagged branches was significantly higher than the percentage of naturally dropped fruits in bagged branches, suggesting that frugivorous birds play an important role in fruit removal of C. franchetii in the study area. Seeds from bird faeces and from manually de‐pulped fruits germinated in higher percentage and faster than seeds from intact fruits. Germination percentage and speed of seeds from manually de‐pulped fruits were significantly higher than those of gut‐passed seeds. These results indicate that T. chiguanco increases and accelerates seed germination of C. franchetii through pulp removal, but not through seed scarification. Overall, our findings indicate that native frugivorous birds facilitate the dispersal and germination success of C. franchetii, likely playing an important role in its invasion throughout the mountainous region of central Argentina.     

Avian fruit consumption and seed dispersal in a temperate Australian woodland

Abstract The effectiveness of avian fruit consumers as seed dispersers of fleshy‐fruited plants was studied in a temperate woodland community. As a consequence of the short and overlapping fruiting phenologies of fleshy‐fruited plant species in temperate regions of Australia, there are very few avian species that are true specialist frugivores. The relative importance of bird species as fruit consumers was investigated, and how their foraging activities, movements and gut treatment of seeds affected dispersal of viable seeds away from the parent plant was examined. Fruit consumption and consumer seed dispersal capacity were assessed in this study through faecal analyses and by testing the viability of seeds that had passed through the gut of avian consumers. Behavioural observations enabled us to determine the consumption rates of, and quantities of fruit consumed by, various bird species and the amount of time spent feeding. Silvereyes (Zosterops lateralis) were the dominant fruit consumers in the community, although 19 bird species were either observed consuming fruit or provided faecal samples that contained fruit. Silvereyes had a high local abundance at the site and more than 90% of silvereyes’faecal samples contained the seeds of fruiting plants (n = 409). Large numbers of fruit were consumed per visit by silvereyes, particularly for Rhagodia parabolica (fragrant saltbush). Silvereyes consumed an average of four R. parabolica fruit per 5 s and up to a maximum 40 fruit per visit. Viability was high for seeds recovered from silvereyes’faeces (R. parabolica, 94.4% viable; Hymenanthera dentata, 100% viable). However, the number of seeds per faecal sample was high for R. parabolica, which may result in density‐dependent seed mortality. Gut passage rate for silvereyes fed R. parabolica fruit in captivity was 31.5 ± 1.9 min. Silvereyes remained at fruiting plants for very short periods (average 50‐60 s) and in most cases moved away from the parent plant, primarily toward canopy trees. Given the short visit duration of silvereyes, individuals would have left the parent plant well before seeds passed through the gut. Rhagodia parabolica fruit was consumed by a large number of bird species in the community, including species often thought of as exclusively insectivorous or nectarivorous. These species are likely to disperse viable R. parabolica seeds into microhabitats different from those visited by silvereyes.     

Linking frugivore activity to early recruitment of a bird dispersed tree,Eugenia umbelliflora (Myrtaceae) in the Atlantic rainforest

Seed dispersal by animals is a complex process involving several distinct stages: fruit removal by frugivores, seed delivery in different microhabitats, seed germination, seedling establishment, and adult recruitment. Nevertheless, studies conducted until now have provided scarce information concerning the sequence of stages in a plant's life cycle in its entirety. The main objective of this study was to evaluate the immediate consequences of frugivore activity for Eugenia umbelliflora (Myrtaceae) early recruitment by measuring the relative importance of each fruit‐eating bird species on the establishment of new seedlings in scrub and low restinga vegetation in the Atlantic rainforest, Brazil. We conducted focal tree observations on E. umbelliflora trees recording birds' feeding behaviour and post‐feeding movements. We also recorded the fate of dispersed seeds in scrub and low restinga vegetation. We recorded 17 bird species interacting with fruits in 55 h of observation. Only 30% of the handled fruits were successfully removed. From 108 post flight movements of exit from the fruiting trees, 30.6% were to scrub and 69.4% to low restinga forest. Proportion of seed germination was higher in low restinga than in the scrub vegetation. Incorporating the probabilities of seeds' removal, deposition, and germination in both sites, we found that the relative importance of each frugivorous bird as seed dispersers varies largely among species. Turdus amaurochalinus and Turdus rufiventris were the best dispersers, together representing almost 12% probability of seed germination following removal. Our results show the importance of assessing the overall consequence of seed dispersal within the framework of disperser effectiveness, providing a more comprehensive and realistic evaluation of the relative importance of different seed dispersers on plant population dynamics.     

Effect of parasitoids,seed‐predators and ant‐mutualists on fruiting success and germination of Acacia drepanolobium in Kenya

Acacia drepanolobium is an obligate ant‐plant that bears dehiscent pods exploited by predispersal seed‐predators and parasitoids. Fruit set and seed germination in relation to ant‐association, bruchid and parasitoid infestation were investigated in a large‐scale multi‐year study in Kenya. Ant‐association had a significant impact on the overall numbers of fruiting trees. 94.8% of Crematogaster mimosae‐occupied trees, 25.6% of C. nigriceps‐occupied trees, and 82.2% of Tetraponera penzigi‐occupied trees set fruit. Within each tree, ant‐association had no significant impact on the amount of seed produced: C. mimosae‐occupied trees produced 47.5 seeds per branch, C. nigriceps‐occupied trees 44.9 seeds per branch and T. penzigi‐occupied trees 38.3 seeds per branch. Ant‐association did not limit seed‐predators or vary significantly by ant. Seeds infested by bruchids germinated in significantly lower proportions (6.2%) compared to uninfested seed (78.6%). Bruchid‐infested seed is also exploited by parasitoid wasps. Parasitoids appear to have a moderate but significant ‘rescue’ effect on bruchid‐infested seed with 18.4% of parasitoid‐infested seed germinating. Stable isotopes (δ15N) revealed the trophic structure of the seed‐associated insects, showing clearly that bruchids are seed predators and the parasitoid wasp Dinarmus magnus exploits the dominant Bruchidius sp.     

The Influence of Temporal Changes in Fruit Availability on Diet Composition and Seed Handling in Blue Monkeys (Cercopithecus mitis doggetti)1

We investigated the relation between temporally varying resources, diet composition, and seed-handling behaviors in a group of blue monkeys (Cercopithecus mitis doggetti) in a tropical montane forest of Rwanda. Changes in diet composition were related to concurrent phenological studies of fruit-producing trees, and density and abundance of tree resources within the monkey's home range. Fruit composed nearly 50 percent of the diet. Over 50 percent of the fruits eaten had juicy fleshy pulp. Observations of seed handling behavior provided insights into the role of these animals as potential seed dispersal agents. The monkeys moved the seeds of 29 species out of parent canopies by defecating seeds intact and by potentially carrying seeds in cheek pouches and dropping them later. Seeds of 18 species were found intact in fecal piles. Our study showed community-level phenology patterns did not indicate a decrease in fruit availability during the study period, but an analysis of the preferred fruits consumed by the monkeys showed distinct periods of low fruit availability. The study period included two dry seasons; only one of these produced a period of fruit scarcity for the animals. The animals employed different strategies during times of preferred fruit scarcity. They increased consumption of leaves and other fleshy fruits, and diet diversity increased, or became mainly seed predators and diet diversity decreased. The variable responses of these monkeys to changes in food availability highlights their dietary plasticity and imposes significant variations in their role as potential seed dispersers.     

Spatial–temporal distribution of ornithochorous seeds from an Elaeagnus umbellata community dominating a riparian habitat

The dispersal efficiency and potential distribution of ornithochorous seeds of Elaeagnus umbellata in a riparian habitat were evaluated to clarify this species' establishment site in relation to the disturbance regime of the floodplain. Fruit removal by avian frugivores was monitored using fruit bags, and the spatial distribution of excreted seeds was quantified by seed traps set randomly on a gravel bar as an isolated seed source in the Yoshino River throughout an autumn fruiting season. Although more than 45% of the fruits remained on the twigs in the fruit bags, almost all fruits on the control twigs without fruit bags were exploited by the beginning of January. The fruit removal rate and seed dispersal distance were positively correlated with an increase in wintering bird species and their abundance. Intact bird‐dispersed seeds of E. umbellata were trapped within a 400‐m range and damaged seeds were limited to traps set within 50 m from the seed source. Frugivore behavior, such as feces excretion on rocks near water drinking sites and perching on surrounding woodland, greatly influenced the spatial and temporal dispersal pattern of the seed rain. In the present study, the avian frugivores showed upstream seed dispersal; thus, in years with stochastic autumnal floods, secondary dispersal via hydrochory downstream may be facilitated. The intensive seed dispersal in E. umbellata indicates that the present distribution of parent trees in the restricted elevation range of the gravel bars is the result of survival through disturbance, rather than seed dispersal limitation.     

Seedling Ecology and Restoration of Blackbrush (Coleogyne ramosissima) in the Mojave Desert,United States

Increases in fire frequency are disrupting many ecological communities not historically subjected to fire. In the southwestern United States, the blackbrush (Coleogyne ramosissima) community is among the most threatened, often replaced by invasive annual grasses after fire. This long‐lived shrub is vulnerable because it recruits sporadically, partially due to mast seeding and the absence of a seed bank. The goal of this study was to evaluate if shrub restoration can be enhanced by identifying and ameliorating recruitment limitations. Specifically, we tested the effect of encapsulating seeds in predation‐deterring “seed balls.” We also tested the effects of nurse plants and mammalian exclusion cages on seedling emergence, growth, and survivorship. These experiments were conducted in a full‐factorial design across three sites differing in elevation. Over 2 years, 13% of all planted seeds emerged and the effect of seed balls was overwhelmingly negative because of low emergence. Nurse plants had overall positive effects at Low Elevation, but negative effects at Mid‐ and High Elevation. Emergence and survival were highest in caged plots everywhere, and effect sizes increased with elevation. Interactions between the cage and the nurse plant treatments indicated that nurse plants tended to attract mammalian predators, lowering emergence and seedling survivorship, particularly at higher elevations. Findings conform to the stress‐gradient hypothesis in that interactions among seedlings and mature plants shifted from facilitation to competition as environmental stress decreased with increasing elevation, suggesting that they are transferable to ecologically similar communities elsewhere. Knowledge of site‐specific recruitment limitations can help minimize ineffective restoration efforts.     

Seed rain of fleshy and dry propagules in different habitats in the temperate rainforests of Chiloé Island,Chile

In temperate rainforests on Chiloé Island in southern Chile (42°S), most canopy trees bear fleshy, avian‐dispersed propagules, whereas emergent tree species have dry, wind‐borne propagules. In the present study, the following hypothesis was tested: regardless of species, fleshy propagules are deposited in greater numbers in canopy gaps and in forest margins and hence have a more heterogeneous seed shadow than wind‐dispersed propagules. To test this hypothesis, the seed rains of these two types of propagules were compared in the following forest habitats: (i) tree‐fall gaps (edges and centre); (ii) forest margins with adjacent pastures; and (iii) under closed canopy (forest interior). Seed collectors (30‐cm diameter) were placed in two (15 and 100 ha) remnant forest patches (n = 60–100 seed collectors per patch) distributed in the four habitats. Seeds were retrieved monthly from each collector during two reproductive seasons (1996, 1997). In both years, the seed rain was numerically dominated by two species with dry propagules (Laureliopsis philippiana and Nothofagus nitida) and three species with fleshy fruits (Drimys winteri, Amomyrtus luma, and Amomyrtus meli). The seed shadows of the two species with dry, wind‐dispersed seeds differed markedly. Seeds of L. philippiana were deposited predominantly in canopy openings, whereas N. nitida seeds fell almost entirely in the forest interior. The fleshy‐fruited species, Drimys and Amomyrtus spp., had similar seed deposition patterns in the various habitats studied, but the between‐year differences in seed rain were greater in Drimys winteri than in Amomyrtus spp. Although no more than 10% of fleshy‐fruited propagules reached the margins of the patch, approximately 7% of these were carried there by birds. Every year, canopy gaps (pooling data from edges and centres) concentrated approximately 60% of the total seed rain of both propagule types in both forest patches. Forest margins received less than 20% of the total seed rain, which was largely dominated by fleshy‐fruited species. Seed shadows were a species‐specific attribute rather than a trait associated with propagule type and dispersal mode.     

Insects remove more seeds than mammals in first‐year prairie restorations

Seed sowing is a common early step in restoration, but seed consumers can impede plant establishment and alter community structure. Moreover, seed consumers vary in feeding behaviors and the relative importance of different seed consumer groups during restoration are not well understood. At 12 first‐year prairie restorations in Michigan, we studied seed predation using seed removal trays to ask: What is the relative magnitude of seed removal by insects and mammals? Do seed removal rates change over the growing season? Do habitat edges influence seed removal? At what rates are 10 prairie plant species' seeds removed by mammals and insects? Seed removal depended on consumer type, time of year, and seed species. Insects accounted for the majority of seed removal, contrary to previous research in similar systems. In May, insects removed 1.8 times more seeds than mammals, while in August, they removed 5.1 times more. There was greater seed removal in August. During May 28% of seeds were removed, compared to 54% of seeds removed during August, an increase driven by insects. Edge proximity did not influence seed removal. Certain seed species were removed more than others. For example, Lespedeza capitata (round‐headed bush clover) was always removed at high rates, whereas Coreopsis lanceolata (lance‐leaved coreopsis) and Andropogon gerardii (big bluestem) were always removed at low rates. Mammals and insects showed different preferences for several species. This research suggests a prominent role of seed predation, particularly by insects, for early prairie restoration dynamics, with influences varying temporally and among species.     

Gut passage effect of the introduced red‐whiskered bulbul (Pycnonotus jocosus) on germination of invasive plant species in Mauritius

In Mauritius, many of the worst invasive plant species have fleshy fruits and rely on animals for dispersal. The introduced red‐whiskered bulbul (Pycnonotus jocosus) feeds on many fleshy‐fruited species, and often moves from invaded and degraded habitats into higher quality native forests, thus potentially acting as a mediator of continued plant invasion into these areas. Furthermore, gut passage may influence seed germination. To investigate this, we fed fleshy fruits of two invasive plant species, Ligustrum robustum and Clidemia hirta, to red‐whiskered bulbuls. Gut passage times of seeds were recorded. Gut‐passed seeds were sown and their germination rate and germination success compared with that of hand‐cleaned seeds, as well as that of seeds in whole fruits. Gut passage and hand‐cleaning had significant positive effects on germination of both species. Gut‐passed seeds of both C. hirta and L. robustum germinated faster than hand‐cleaned seeds. However, for L. robustum, this was only true when compared with hand‐cleaned seeds with intact endocarp; when compared with hand‐cleaned seeds without endocarp, there was no difference. For overall germination success, there was a positive effect of gut passage for C. hirta, but not for L. robustum. For both C. hirta and L. robustum, no seeds in intact fruits geminated, suggesting that removal of pulp is essential for germination. Our results suggest that, first, the initial invasion of native forests in Mauritius may not have happened so rapidly without efficient avian seed dispersers like the red‐whiskered bulbul. Second, the bulbul is likely to be a major factor in the continued re‐invasion of C. hirta and L. robustum into weeded and restored conservation management areas.