Rodent seed predation: effects on seed survival, recruitment, abundance, and dispersion of bird-dispersed tropical trees

Tropical tree species vary widely in their pattern of spatial dispersion. We focus on how seed predation may modify seed deposition patterns and affect the abundance and dispersion of adult trees in a tropical forest in India. Using plots across a range of seed densities, we examined whether seed predation levels by terrestrial rodents varied across six large-seeded, bird-dispersed tree species. Since inter-specific variation in density-dependent seed mortality may have downstream effects on recruitment and adult tree stages, we determined recruitment patterns close to and away from parent trees, along with adult tree abundance and dispersion patterns. Four species (Canarium resiniferum, Dysoxylum binectariferum, Horsfieldia kingii, and Prunus ceylanica) showed high predation levels (78.5-98.7%) and increased mortality with increasing seed density, while two species, Chisocheton cumingianus and Polyalthia simiarum, showed significantly lower seed predation levels and weak density-dependent mortality. The latter two species also had the highest recruitment near parent trees, with most abundant and aggregated adults. The four species that had high seed mortality had low recruitment under parent trees, were rare, and had more spaced adult tree dispersion. Biotic dispersal may be vital for species that suffer density-dependent mortality factors under parent trees. In tropical forests where large vertebrate seed dispersers but not seed predators are hunted, differences in seed vulnerability to rodent seed predation and density-dependent mortality can affect forest structure and composition.     

Rodent seed predation and seedling recruitment in mesic grassland

Seedling recruitment of two grasses (Arrhenatherum elatius and Festuca rubra) and two herbs (Centaurea nigra and Rumex acetosa) was measured in areas with and without rodents to which seeds of each species were sown at three seed densities (1000, 10,000 and 50,000 seeds m⁻²) in two seasons (spring and autumn 1995). Seed removal was measured for 10-day periods and the fate of seedlings was followed for 15 months after sowing. The proportion of seed removed ranged from 6 to 85% and increased with increasing seed density for each species. Rodents had no effect on seedling emergence or survival in the spring sowing. In the autumn sowing, rodents reduced seedling emergence of all four species sown at 1000 and 10,000 seeds m⁻² but had no impact at 50,000 seeds m⁻², presumably because of microsite limitation. We suggest the difference between spring and autumn arose because emergence was seed limited in autumn but microsite limited in spring; microsite availability was higher in autumn because a summer drought killed plants, reduced plant biomass and opened up the sward. Fifteen months after the autumn sowing, fewer A. elatius and C. nigra seedlings survived on plots exposed to rodents. This result reflected not only the reduced seedling emergence but also increased seedling mortality (seedling herbivory) in sites exposed to rodents. In contrast, F. rubra and R.acteosa showed density-dependent seedling survival which compensated for initial differences in seedling emergence, so that no effect of rodents remained after 15 months. The results suggest that rodent seed predation and seedling herbivory exert strong effects on seedling recruitment of A.elatius and C. nigra when recruitment conditions are favourable (conditions that lead to high microsite availability) and may contribute to both species being maintained at low densities in the grassland. The results also demonstrate that highly significant impacts of rodent seed predation at the seedling emergence stage can disappear by the time of plant maturation. Received: 2 March 1998 / Accepted: 28 September 1998     

A meta-analysis of biotic resistance to exotic plant invasions

Biotic resistance describes the ability of resident species in a community to reduce the success of exotic invasions. Although resistance is a well‐accepted phenomenon, less clear are the processes that contribute most to it, and whether those processes are strong enough to completely repel invaders. Current perceptions of strong, competition‐driven biotic resistance stem from classic ecological theory, Elton's formulation of ecological resistance, and the general acceptance of the enemies‐release hypothesis. We conducted a meta‐analysis of the plant invasions literature to quantify the contribution of resident competitors, diversity, herbivores and soil fungal communities to biotic resistance. Results indicated large negative effects of all factors except fungal communities on invader establishment and performance. Contrary to predictions derived from the natural enemies hypothesis, resident herbivores reduced invasion success as effectively as resident competitors. Although biotic resistance significantly reduced the establishment of individual invaders, we found little evidence that species interactions completely repelled invasions. We conclude that ecological interactions rarely enable communities to resist invasion, but instead constrain the abundance of invasive species once they have successfully established.     

Sex-biased seed predation and the maintenance of females in a gynodioecious plant

We investigated genetic and ecological factors contributing to the maintenance of females in populations of the gynodioecious plant, Sidalcea hendersonii (Malvaceae). Our crossing experiments indicated that male sterility is controlled by a dominant nuclear allele. Nuclear determination of sex theoretically requires much higher female fitness to account for the high frequency of female plants often observed during surveys of S. hendersonii populations. Females were, in fact, shown to have higher fitness in an experimental population, producing more viable seed and surviving offspring than hermaphrodite plants. In natural populations, however, differences in viable seed production were not evident unless seed predation was considered. Sex-biased seed predation appears to play an important role in the maintenance of gynodioecy in S. hendersonii. In populations with high female frequencies, weevil larvae destroyed significantly more seeds from hermaphrodite plants than females, substantially reducing their seed production. To our knowledge, this study provides the first evidence that sex-biased predation may be responsible for high female frequencies in natural populations of a gynodioecious species.     

Evidence of biotic resistance to exotic plant invasion in degraded Bornean forests

Intact tropical forests are generally considered to be resistant to invasions by exotic species, although the shrub Clidemia hirta (Melastomataceae) is highly invasive in tropical forests outside its native range. Release from natural enemies (e.g., herbivores and pathogens) contributes to C. hirta invasion success where native melastomes are absent, and here we examine the role of enemies when C. hirta co-occurs with native Melastomataceae species and associated herbivores and pathogens. We study 21 forest sites within agricultural landscapes in Sabah, Malaysian Borneo, recording herbivory rates in C. hirta and related native Melastoma spp. plants along two 100-m transects per site that varied in canopy cover. Overall, we found evidence of enemy release; C. hirta had significantly lower herbivory (median occurrence of herbivory per plant = 79% of leaves per plant; median intensity of herbivory per leaf = 6% of leaf area) than native melastomes (93% and 20%, respectively). Herbivory on C. hirta increased when closer to native Melastoma plants with high herbivory damage, and in more shaded locations, and was associated with fewer reproductive organs on C. hirta. This suggests host-sharing by specialist Melastomataceae herbivores is occurring and may explain why invasion success of C. hirta is lower on Borneo than at locations without related native species present. Thus, natural enemy populations may provide a “biological control service” to suppress invasions of exotic species (i.e., biotic resistance). However, lower herbivory pressures in more open canopy locations may make highly degraded forests within these landscapes more susceptible to invasion.     

Rodent seed predation promotes differential recruitment among bird-dispersed trees in temperate secondary forests

We investigated the role of seed predation by rodents in the recruitment of the fleshy-fruited trees Taxus baccata, Ilex aquifolium and Crataegus monogyna in temperate secondary forests in NW Spain. We measured the densities of dispersed seeds, early emerged seedlings, established recruits and adults, at four sites over a period of 2 years. Seed predation among species was compared by seed removal experiments and analysis of rodent larder-hoards. The three species differed markedly in local regeneration patterns. The rank order in the seed rain following decreasing seed density was Ilex, Taxus and Crataegus. However, Crataegus established 3.3 times more seedlings than Taxus. For all species, there was a positive linear relationship between the density of emerged seedlings and seed density, suggesting that recruitment was seed- rather than microsite-limited. A consistent pattern of seed selection among species was exerted by rodents, which preferred Taxus and, secondarily, Ilex seeds to Crataegus seeds. Predation ranking was the inverse of that of seed protection against predators, measured as the mass of woody coat per mass unit of the edible fraction. Recruitment potential, evaluated as the ratio of seedlings to seeds, was negatively related to seed predation, with the rank order Crataegus > Ilex > Taxus. The selective early recruitment limitation exerted by predation may have a demographic effect in the long term, as judged by the positive relationship between early seedling emergence and the density of established recruits. By modulating the pre-emptive competition for seed safe sites, rodents may preclude the progressive exclusion of species that produce low numbers of seeds (i.e. Crataegus) by those dominant in seed number (i.e. Ilex, Taxus), or at least foster the evenness for site occupation among seedlings of different species.     

Post-dispersal seed predation: consequences for plant demography and evolution

Post-dispersal seed predation is only one of many factors underlying plant demography and evolution. Nevertheless, the generalist feeding habits of many post-dispersal seed predators and the limited ability of plants either to compensate for or to respond to post-dispersal seed losses directly suggest that post-dispersal seed predation may have a considerable impact on plant populations. Seed predators probably have little direct influence on the demography of plants that regenerate exclusively by vegetative means or are buffered by a large active seed bank, but such species are only a minority in most plant communities.In general, ants are significant post-dispersal seed predators in arid and semi-arid ecosystems while they act mainly as seed dispersers rather than as predators in temperate ecosystems. Although studies have probably underestimated the importance of invertebrates and birds as seed predators, rodents appear to have greater potential to influence seed dynamics, and are particularly important in temperate ecosystems. For example, production of mast seed crops is more effective at satiating specialist invertebrate seed predators than generalist vertebrates, and recruitment may be limited by post-dispersal seed predation even during mast years.Both spatial variation in post-dispersal seed predation and differences in predation between species are important elements which facilitate the coexistence of different plant species. Where microsites are limiting, selective post-dispersal seed predators can influence pre-emptive competition for these microsites. Seed size determines the extent of density-dependent predation and the exploitation of buried seed. This suggests that post-dispersal seed predators may also play a role in the evolution of seed characteristics. However, conclusions regarding the ecological and evolutionary impact of post-dispersal seed predators will remain speculative without a more substantial empirical base.     

Plant community diversity and native plant abundance decline with increasing abundance of an exotic annual grass

Exotic plants are generally considered a serious problem in wildlands around the globe. However, some argue that the impacts of exotic plants have been exaggerated and that biodiversity and other important plant community characteristics are commonly improved with invasion. Thus, disagreement exists among ecologists as to the relationship of exotic plants with biodiversity and native plant communities. A better understanding of the relationships between exotic plants and native plant communities is needed to improve funding allocation and legislation regarding exotic plants, and justify and prioritize invasion management. To evaluate these relationships, 65 shrub–bunchgrass plant communities with varying densities of an exotic annual grass, Taeniatherum caput-medusae (L.) Nevski (medusahead), were sampled across 160,000 ha in southeastern Oregon, United States. Environmental factors were generally not correlated with plant community characteristics when exotic annual grass density was included in models. Plant diversity and species richness were negatively correlated with exotic annual grass density. Exotic annual grass density explained 62% of the variation in plant diversity. All native plant functional groups, except annual forbs, exhibited a negative relationship with T. caput-medusae. The results of this study suggest that T. caput-medusae invasions probably have substantial negative impacts on biodiversity and native plant communities. The strength of the relationships between plant community characteristics and T. caput-medusae density suggests that some exotic plants are a major force of change in plant communities and subsequently threaten ecosystem functions and processes. However, experimental studies are needed to fully substantiate that annual grass invasion is the cause of these observed correlations.     

Post-dispersal seed predation of three grassland species in a plant diversity experiment

Aims Post-dispersal seed predation is an important ecosystem process because it can influence the seed's fate after the initial dispersal from the mother plant and subsequently transform communities. Even at small scales, post-dispersal seed predation can vary greatly depending on seed identity, granivorous taxa or microhabitat structure. However, little is known about the role of plant species richness and functional group richness in post-dispersal seed predation. The overall aim of this study was to test whether increasing plant species richness or plant functional group richness affects the rate and variability of post-dispersal seed predation. We additionally investigated the influence of vegetation structure and seed species identity on the rate and variability of post-dispersal seed predation and whether the influence of different granivorous taxa changed with increasing plant species richness.Methods We conducted seed removal experiments along a long-term experimental plant diversity gradient, comprising plots with monocultures to 60 species mixtures of common grassland species in Jena, Germany, in August 2011. We studied seeds of Onobrychis viciifolia, Pastinaca sativa and Trifolium pratense in exclusion experiments (seed cafeterias), an experimental setup that allowed access either for arthropods or slugs or for all granivorous taxa. Traditionally, seeds removed from seed cafeterias were classified as consumed but we used traceable fluorescent-coloured seeds to obtain more accurate predation rates by subtracting recovered seeds from overall removed seeds. The effect of multiple vegetation variables on mean and variability of seed predation rates was analysed using generalized mixed-effect models and linear regressions, respectively.Important findings Rates of recovered seeds were low but contributed to significant differences between seed predation rates and removal rates of seeds in some treatments. Seed predation rates were not directly correlated with increasing plant species richness or plant functional group richness but were influenced byseed species identity and granivorous taxa. Vegetation variables such as vegetation height and cover were significantly associated with seed predation rates. Depending on the seed species and/or the granivorous taxa, different vegetation variables correlated with seed predation rates. Our results indicate that effects of plant functional group richness and multiple vegetation variables on the magnitude of post-dispersal seed predation varied with seed identity and seed predator taxa. A direct effect of plant species and plant functional group richness could be shown on the variability of post-dispersal seed predation for some seed species and their respective predators. Thus, the changes in magnitude of post-dispersal seed predation with increasing plant species richness could potentially impact the fitness of some plant species and thereby influence plant community structure.     

Small mammal abundance and seed predation across boundaries in a restored‐grazed woodland interface

Passive restoration is an effective tool for the maintenance and conservation of biodiversity. Often areas in recovery are immersed in a matrix of land uses, in which the expansion and intensification of human activities exert new visible pressures at their boundaries. The degree of connectivity between these areas and their peripheral lands can be analyzed by mobile link species, organisms that actively move in the landscape by connecting areas to one another through their functional roles. We focus our design on the interface generated by the long‐term restoration area and surrounding grazing lands. We analyze the changes on boundary structure, small mammal abundance, and on the function of native seed dispersal by these vertebrate species. We captured small mammals and determined seed removal of Prosopis flexuosa at three distances inside and outside a fence that delineates passively restored and currently grazed areas. Our results indicate that small rodents find more suitable habitats at the site under restoration than in grazing lands. The restored‐grazing interface shows a decrease in small mammal abundance from the protected area to the grazed lands. From a functional perspective, an increase in small mammal abundance results in an increase in their seed removal activity with implications for seed fate, because the long‐term recovery of vegetation could enhance seed predation on a native tree species.     

Rodent seed predation on tree invader species in grassland habitats of the inland Pampa

Woody plant invasion in grassland ecosystems is a worldwide phenomenon, and biotic interactions as competition and predation have been invoked as a possible barrier to woody encroachment in many ecosystems. We evaluated the role of rodents as seed predators in Pampean grasslands, and we assessed the differences in removal by rodents between one native species, Prosopis caldenia (Caldén) and one exotic species, Gleditisia triacanthos (Honey locust). The experiment was conducted at different phases of the rodent population cycle in two grassland communities, a remnant of a native grassland and a post agriculture grassland (old field). The amount of seed loss caused by predation was estimated by a bait-removal experiment in foraging stations. We estimated the frequency of foraging stations with consumption, the overall amount of seed predation and the individual rate of seed predation. The total amount of seed removal and the individual rate of seed removal were higher for P. caldenia than for G. triacanthos, in the native grassland than in the old field, and in autumn when rodent density was maximum. Overall, the role of rodents on woody seed removal varied according to the plant species and depending on the local conditions that vary through time and space.     

Rodent seed predation in cereal crop areas of central Spain: effects of physiognomy, food availability, and predation risk

Patterns of rodent seed predation in extensive cereal croplands of central Spain were investigated by measuring rodent predation rates on artificial seed patches, together with seed availability, rodent abundance and habitat physiognomy Sampling was designed to cope with the spatial and temporal variability of factors potentially affecting rodent activity (natural and man-made habitat features, season and moonlight) Rodent predation was unpredictably distributed in summer In winter, rodents foraged on average in the patches least used by man. moonlight, however, did alter this pattern, so that rodent predation was closely associated with shrubs in the bright nights and evenly distributed in the dark ones I conclude that the need of stable underground refuges and predation risk in the winter period seemed to be the mam factors constraining rodent foraging activity in the man-made ecosystem studied     

An exotic magnet plant alters pollinator abundance and behavior: a field test with a native mistletoe

Exotic species can threaten biodiversity by disrupting ecological interactions among native species. Highly-attractive exotic species can exert a ‘magnet effect’ by attracting native pollinators, which may have either competitive or facilitative effects on co-flowering native plants. However, those effects may be context-dependent. We used a mistletoe-hummingbird pollination system in the Valdivian rainforest (southern Chile) to test whether the exotic tree Eucalyptus globulus (a highly attractive species to pollinators) acts as a magnet species, affecting the co-flowering native mistletoe. We compared hummingbird abundance, visitation rates, and activity patterns between native forest and abandoned E. globulus plantations. We found that hummingbirds were more abundant and visited more flowers at the plantation irrespective of E. globulus flowering. We observed a significant change of pollinator activity at the native habitat during E. globulus flowering, as hummingbirds visited mistletoe flowers more frequently early in the morning at the plantations and in the afternoon at the native forests. Our results showed that E. globulus acts as an exotic magnet species and can alter pollinator abundance and behavior. Our findings demonstrate the importance of considering local- and landscape-scale processes to understand the effects of magnet species on native plants and suggest that magnet species may influence even highly-attractive plants.

     

Impact of habitat structure and fruit abundance on avian seed dispersal and fruit predation

So far, it is poorly understood how differential responses of avian seed dispersers and fruit predators to changes in habitat structure and fruit abundance along land-use gradients may translate into consequences for the seed dispersal of associated plants. We selected a gradient of habitat modification (forest, semi-natural, and rural habitat) characterized by decreasing tree cover and a high variation in local fruit availability. Along this gradient we quantified fruit removal by avian seed dispersers and fruit predators from 18 Sorbus aucuparia trees. We analyzed the relative importance of tree cover and fruit abundance in explaining species richness, abundance and fruit removal rates of both guilds from S. aucuparia trees. Species richness and abundance of seed dispersers decreased with decreasing tree cover, whereas fruit removal by seed dispersers decreased with decreasing fruit abundance independent of tree cover. Both variables had no effect on species richness, abundance and fruit removal by fruit predators. Consequently, seed dispersers dominated relative fruit removal in fruit-rich sites but the dispersal/predation ratio shifted in favor of predation in fruit-poor habitat patches. Our study demonstrates that variation in local habitat structure and fruit abundance can cause guild-specific responses. Such responses may result in a shift in fruit removal regimes and might affect the dispersal ability of dependent fruiting plants. Future studies should aim at possible consequences for plant recruitment and guild-specific responses of frugivores to disturbance gradients on the level of entire plant–frugivore associations.     

Abiotic and biotic factors influencing the winter distribution of predatory insects

Summary Various environmental factors were investigated to analyse those involved in successful overwintering and possibly overwintering site selection for Tachyporus hypnorum and Demetrias atricapillus, both important coleopteran predators of cereal aphids. The results of the study indicated food supply to be important for both predator species during the winter period, although the role of biotic factors in site selection in the autumn could not be clearly demonstrated. The winter distribution of the two species could, however, be explained well in terms of abiotic factors. It is suggested that these and other similar predator species have well-defined overwintering requirements and that these can be exploited in the management of field boundary habitats.     

On the distribution of plant abundance data

Plant abundance data are often analysed using standard statistical procedures without considering their distributional features and the underlying ecological processes. However, plant abundance data, e.g. when measured in biodiversity monitoring programs, are often sampled using a hierarchical sampling procedure, and since plant abundance data in a hierarchical sampling procedure are typically both zero-inflated and over-dispersed, the use of a standard statistical procedure is sub-optimal and not the best possible practice in the modelling of plant abundance data. Two distributions (the zero-inflated generalised binomial distribution and the zero-inflated bounded beta distribution) are suggested as possible distributions for analysing either discrete, continuous, or ordinal hierarchically sampled plant cover data.