Mammalian predator scent, vegetation cover and tree seedling predation by meadow voles

Herbivores are thought to respond to the increased risk of attack by predators during foraging activities by concentrating feeding in safe habitats and by reducing feeding in the presence of predators. We tested these hypotheses by comparing tree seedling predation by meadow voles within large outdoor enclosures treated either with scent of large mammalian predators (red fox, bobcat, coyote) or a control scent (vinegar). In addition, we compared the distribution of voles in relation to naturally occurring variation in vegetation cover and the tendency of voles to attack tree seedlings planted in small patches with cover manipulation (intact, reduced or removed cover). Predator scent did not affect the rate or spatial distribution of tree seedling predation by voles, nor did it affect giving up densities (a surrogate of patch quitting harvest rate), survival rates, body size or habitat distribution of voles. In both predator scent and vinegar treatments voles preferred abundant vegetation providing good cover, which was also the site of almost all tree seedling predation. We conclude that large mammalian predator scent does not influence the perception by voles of the general safety of habitat, which is more strongly affected by the presence of cover.     

啮齿类取食的物种偏好与时空格局

通过强烈消耗土壤种子库,动物取食种子对植物种群更新和群落动态产生深远的影响。一般认为种子被食概率的空间格局取决于种子密度和离母树的距离,而环境（如地形）异质性的影响则一直没有得到足够的关注,与此相关的机制及其影响程度亦不清楚。研究设计在野外埋放种子以模拟种子扩散后的情形,监测啮齿类对种子的取食,以检验种子取食受埋藏生境、时间及动物对种子种类的偏好等因素的影响。结果表明,经过1a实验,8种落叶阔叶树种子的累计取食概率为0～48．25％,平均值为20％;山顶部位的取食概率大致是其它部位概率的3倍;埋放在凋落物层中的种子被食概率大约为埋放在土壤层中概率的2倍。利用logistic回归模型进行统计分析表明,种子被食概率变化的45％可以被上述因素解释。其中,物种偏好是影响种子被取食概率的首要因素,其后依次是地形、埋藏时间和深度。啮齿类明显喜好较大的种子;其取食行为在山脊部位明显较其它部位更频繁和剧烈;对埋藏种子的取食从3月份开始加剧,到7月份以后平息下来。种子埋放深度对啮齿类的取食概率有显著影响。     

Seed predation and seedling recruitment in plants: the effect of the distance between parents

We present a graphic model that explores the effect of distance between parent plants on seed predation and seedling recruitment. Based on the assumption that distance between parents may affect the shape of the seed shadow, the model predicts that seed predators may affect seedling recruitment curves under isolated plants but they are unable to affect these curves under close parent plants. The predictions of the model are tested experimentally in Cryptocarya alba (Lauraceae), a common tree of the Mediterranean forest, Central Chile. Results show that predictions are not met under isolated parent plants. Although seed density decreases significantly away from parent plants, this effect is not relevant for seed predation and seedling recruitment. The biotic/abiotic contrast existing under the canopy vs outside the canopy, plus the shade-tolerance of this tree, better explains the seedling recruitment observed under isolated parent plants. Nevertheless, the predictions of the model are corroborated under close parent plants. Seed shadows overlap to the extent that they generate a homogeneous seed distribution, homogeneous seed predation and homogeneous seedling recruitment as well. We discuss the implications of the model in terms of the spatial pattern of seedlings and the benefits of dispersal from isolated and close parent plants.     

Post‐dispersal seed predation and its relations with seed traits: a thirty‐species‐comparative study

Post‐dispersal seed predation is a key process determining the variability in seed survival in forests, where most seeds are handled by rodents. Seed predation is thought to affect seedling regeneration, colonization ability and spatial distribution of plants. Basic seed traits are the essential factors affecting rodent foraging preferences and thus seed survival and seedling recruitment. Many studies have discussed several seed traits and their effects upon seed predation by rodents. However, the results of those previous studies are usually equivocal, likely because few seed traits and/or plant species tend to be incorporated into these studies. In order to elucidate the relationships between seed predation and seed traits, we surveyed the predation of 48 600 seeds in a natural pine forest, belonging to 30 species, for three consecutive years. The results demonstrated that: (i) seed size and seed coat hardness did not significantly affect seed predation; (ii) total phenolics had a negative effect upon seed predation; (iii) positive effects of nitrogen content upon seed predation were found. From our study, it seems that the better strategy to prevent heavy predation is for plants to produce seeds with higher total phenolics content rather than physical defenses (i.e. hard seed coat) or larger seeds. Additionally, rodent foraging preference may depend more on Nitrogen content than other nutrient content of seeds.     

Effects of stoat's presence and auditory cues indicating its presence on tree seedling predation by meadow voles

Predators may control the impact of herbivores on their plant resources by 1) decreasing herbivore numbers, 2) imposing predation risk affecting foraging behavior. The goal of the present study was to examine the effects of a predator and auditory cues indicating its presence on the rate of tree seedling (Acer rubrum, Betula lenta) consumption by meadow voles (Microtus pennsylvanicus). The first of our experiments involved introduction of a stoat (Mustela erminea) into an enclosed vole population and the second a playback of recordings of vole distress calls, movements of a stoat and its vocalizations. In both experiments we manipulated vegetation cover and the availability of food next to the experimental seedlings to assess the effects of microhabitat under the different risk situations. The results of the first unreplicated experiment suggested an increased rate of seedling predation in the presence of the stoat. Consistent with these results, the playback of predator sounds in the second replicated experiment caused an increased rate of seedling predation compared to control plots with no recordings. A mowed circle around a seedling station, representing increased risk of predation on the voles, decreased seedling consumption. This effect was modest in the playback treatment. We suggest the results to be due to displacement behavior by the voles exposed to prolonged risk and conflicting demands of foraging and avoiding predators. Alternatively, as suggested by the model of Lima and Bednekoff, prolonged risk of predation forced the voles to decrease their levels of vigilance during low‐risk playback breaks. The modest inhibitory effect of cover removal on seedling predation in the playback treatment is consistent with this interpretation. The results confirm recent evidence for trophic cascades mediated by behavioral interactions between predator and prey. They are novel in suggesting that the presence of predation risk can increase the inhibitory effects of consumers on their resources.     

Old-field colonization by Daphne gnidium: seedling distribution and spatial dependence at different scales

Abstract. This paper deals with the spatial distribution pattern of the bird-dispersed plant Daphne gnidium in a 10-yr abandoned field under Mediterranean conditions. Colonization of Mediterranean old-fields by bird-dispersed plants is expected to fit a theoretical model in which (1) seed dispersal follows a negative exponential curve with the distance from the seed source and (2) seedlings are better established under perches (nucleation sensu Yarranton & Morrison 1974). However, post-dispersal processes such as seed predation, seed germination and seedling establishment are also spatial-dependent and can lead to spatial autocorrelation in the seedling distribution within an old-field. Results show that both processes in the model (curve of seed dispersal and nucleation) significantly explained the spatial distribution of the seedlings, but some spatial variance remained unexplained. The semivariogram with the statistical residuals of the model detected spatial dependence at small (< 20 m) and large (> 250 m) distance intervals, indicating that some mechanisms with spatial components, apart from the curve of seed dispersal and nucleation under perches, also determined the distribution of seedlings colonizing fields. At scales below 20-m intervals, semivariance increased indicating that similarity between plots is lost when distance increases between them. This pattern may be explained because the favourable micro-environmental conditions for establishment produced under perches could be extended towards neighbour plots where perches were absent. A flat semi-variogram between 20-m and 250-m intervals shows spatial independence along this range. From 250 m on, the semivariance increased again, indicating spatial dependence at coarse-scale. It is possible that the colonization model failed at this scale because different spatial processes to those included in the model (perch presence and distance to the shrubland) could be controlling seed colonization at coarse-scale.     

Consumers limit the abundance and dynamics of a perennial shrub with a seed bank

For nearly 30 years, ecologists have argued that predators of seeds and seedlings seldom have population-level effects on plants with persistent seed banks and density-dependent seedling survival. We parameterized stage-based population models that incorporated density dependence and seed dormancy with data from a 5.5-year experiment that quantified how granivorous mice and herbivorous voles influence bush lupine (Lupinus arboreus) demography. We asked how seed dormancy and density-dependent seedling survival mediate the impacts of these consumers in dune and grassland habitats. In dune habitat, mice reduced analytical lambda (the intrinsic rate of population growth) by 39%, the equilibrium number of aboveground plants by 90%, and the seed bank by 98%; voles had minimal effects. In adjacent grasslands, mice had minimal effects, but seedling herbivory by voles reduced analytical lambda by 15% and reduced both the equilibrium number of aboveground plants and dormant seeds by 63%. A bootstrap analysis demonstrated that these consumer effects were robust to parameter uncertainty. Our results demonstrate that the quantitative strengths of seed dormancy and density-dependent seedling survival--not their mere existence--critically mediate consumer effects. This study suggests that plant population dynamics and distribution may be more strongly influenced by consumers of seeds and seedlings than is currently recognized.     

The effects of habitat manipulation on population distribution and foraging behavior in meadow voles

Individuals, free to choose between different habitat patches, should settle among them such that fitness is equalized. Alternatives to this ideal free distribution result into fitness differences among the patches. The concordance between fitnesses and foraging costs among inhabitants of different quality patches, demonstrated in recent studies, suggests that the mode of habitat selection and the resulting fitness patterns may have important implications to the resource use of a forager and to the survival of its prey. We studied how coarse scale selection between habitat patches of different quality and quitting harvest rate in these patches are related to each other and to fine scale patch use in meadow voles (Microtus pennsylvanicus). To demonstrate these relationships, we manipulated habitat patches within large field enclosures by mowing vegetative cover and adding supplemental food according to a 2×2 factorial design. We tracked vole population densities, collected giving‐up densities (GUDs, a measure of patch quitting harvest rate), and monitored the removal of seeds from lattice grids with 1.5 m intervals (an index of fine‐scale space use) in the manipulated habitat patches. Changes in habitat quality induced changes in habitat use at different spatial scales. In preferred habitats with intact cover, voles were despotic and GUDs were low, but increased with the addition of food. In contrast, voles in less‐preferred mowed habitats settled into an ideal free distribution, GUDs were high and uninfluenced by the addition of food. Seed removal was enhanced by the presence of cover but inhibited by supplemental food. Across all treatments, vole densities and GUDs were strongly correlated making it impossible to separate their effects on seed removal rates. However, this relationship broke down in unmowed habitats, where GUDs rather than vole density primarily influenced seed removal by voles. GUDs and seed removal correlated with predation on tree seedlings formerly planted into the enclosures, demonstrating the mechanisms between coarse‐scale habitat manipulations and community level consequences on a forager's prey.     

Consumer preference for seeds and seedlings of rare species impacts tree diversity at multiple scales

Positive density-dependent seed and seedling predation, where herbivores selectively eat seeds or seedlings of common species, is thought to play a major role in creating and maintaining plant community diversity. However, many herbivores and seed predators are known to exhibit preferences for rare foods, which could lead to negative density-dependent predation. In this study, we first demonstrate the occurrence of increased predation of locally rare tree species by a widespread group of insular seed and seedling predators, land crabs. We then build computer simulations based on these empirical data to examine the effects of such predation on diversity patterns. Simulations show that herbivore preferences for locally rare species are likely to drive scale-dependent effects on plant community diversity: at small scales these foraging patterns decrease plant community diversity via the selective consumption of rare plant species, while at the landscape level they should increase diversity, at least for short periods, by promoting clustered local dominance of a variety of species. Finally, we compared observed patterns of plant diversity at the site to those obtained via computer simulations, and found that diversity patterns generated under simulations were highly consistent with observed diversity patterns. We posit that preference for rare species by herbivores may be prevalent in low- or moderate-diversity systems, and that these effects may help explain diversity patterns across different spatial scales in such ecosystems.     

Dispersion pattern and mortality of seeds and seedlings ofFagus crenata Blume in a cool temperate forest in western Japan

The seed and seedling mortality ofFagus crenata Blume after a mast year (1993) was examined in relation to density and distance from the nearest conspecific adult tree in a mixed conifer-hardwood forest in Ohdaigahara, western Japan. The mortality of fallen seeds during winter amounted to 93.7%, and 79.2% of the current-year seedlings died in the first growing season. The most important factor of death for both seeds and seedlings was predation by vertebrates. The mortality of seeds during winter was positively correlated with sound seed density. The mortality of seedlings was positively correlated with density but not significantly related to the distance from the nearest crown edge of a conspecific adult tree. Mortality patterns varied with stages and spatial scales due to the behavior of predators; it is thus important to investigate the spatial pattern of seeds and seedling mortality at various temporal and spatial scales. After the first growing season, the difference in seedling density between distance classes was not significant at <4m from the nearest adult trees due to density-dependent mortality. However, seedling density was significantly lower in the ≥4 m class than in the <4 m classes.     

Seed predators limit plant recruitment in Neotropical savannas

Despite the well‐documented impacts of consumers on seed abundance the link between seed predation and plant population dynamics remains poorly understood because experimental studies linking patterns of predation with seedling establishment are rare. We used experimental manipulations with six woody plant species to elucidate the effects of seed predator type, habitat, and plant species identity on rates of seed predation and seedling recruitment in the Neotropical savannas known as the Cerrado. We found that seed predation rates are consistently high across a diversity of local habitat types, with important inter‐habitat variation in seed predation for three of the six species used in our experiments. We also found that seed predation has a clear demographic signal – experimentally excluding predators resulted in higher rates of seedling establishment over the course of two seasons. Because the intensity of seed predation varied between species and habitats, it may play a role in structuring local patterns of plant abundance and community composition. Finally, our results lend support to the recent hypothesis that herbivores have major and underappreciated impacts in Neotropical savannas, and that top–down factors can influence the demography of plants in this extensive and biodiversity‐rich biome in previously unexplored ways.     

Tree invasion and establishment in old fields at Hutcheson Memorial Forest

In this review I present results from pattern analysis and field experiments in New Jersey, United States concerning tree invasion and establishment in old fields. Hutcheson Memorial Forest Center (HMF) is the New Jersey study site and a logical choice to anchor a review of this topic. HMF contains the longest and largest old field permanent plot study in the USA including extensive data concerning tree invasion and establishment. HMF has also been the site of considerable old field tree experimental research. In addition, I review other investigations of old field tree invasion and establishment of the eastern deciduous biome and contrast them with HMF. I identify critical questions concerning tree invasion and establishment, indicate what progress has been made towards answering these questions and propose future experimentation. I also synthesize results into a conceptual model and investigate the relative importance of seed and seedling processes in determining tree dynamics in old fields. The idea of a possibility or “window” for tree invasion and establishment and its variation form the basis for this review. For example, the temporal variation of windows is reflected in tree establishment patterns at HMF because wind- and bird-dispersed species invaded one year after cessation of cultivation while mammal-dispersed species were not present until year 12. Although analysis of size classes of individual HMF trees imply that windows close quickly, studies are of too short duration to tell definitely. Spatial variation in invasion and establishment pattern includes exponential decline in seed and stem density with distance from forest edge which deteriorates with time as the old field becomes more structurally complex. Additional spatial heterogeneity occurs when seeds and seedlings of bird-dispersed species clump which conversely becomes more pronounced with time. Many of these HMF patterns are seen in other eastern United States old field sites. When reviewing field experiments from my site and others, I focus on the spatial and temporal variation and the difference between species in the effect of processes on tree seeds and seedlings. I do this because this variation relates directly to the dynamic nature of windows whose quantification is necessary for modeling. Combining my pattern analysis with a survey of variation in seed and seedling mortality caused by different successional processes in old fields, I conclude that seed dispersal, conditions necessary for seed germination and seed and seedling predation are the critical processes controlling and constraining tree invasion and establishment in old fields. Competition and indirect effects of litter and the environment, which modify how these processes affect tree dynamics, are also important. Therefore, processes involving the interaction of species with the environment and with other species form the principal filters on and limitations to tree invasion and establishment and not the non-interactive life-history traits of individual tree species such as growth rate or longevity. The interaction of pattern and process and the affect of many of these processes on old field community structure are also discussed.     

Spatial and temporal patterns of granivorous ant seed predation in patchy cereal crop areas of central Spain

Summary Patterns of granivorous ant seed predation in extensive cereal croplands of central Spain were investigated by measuring seed removal rates on artificial seed patches. Sampling was designed to cope with the seasonal and daily foraging cycle of ant colonies. Simultaneously with removal rates, I measured seed availability, habitat physiognomy at two spatial scales (landscape and microhabitat), weather variables (temperature and rainfall), and distance to the nearest ant nest. Ant seed predation was concentrated on shrublands, and associated with places with high covers of shrubs, chamaephytes and stones. These results were in close agreement with those obtained by analyzing the spatial distribution of granivorous ant nests (Díaz 1991). Moreover, there was a close relationship between seed removal rates and distance to the nearest ant nest, that fitted the predictions of the optimal foraging model developed by Reyes-López (1987). Seasonal and daily patterns of ant foraging activity seemed to depend more on endogenous factors than on environmental variation. I conclude that ants were not able to track the spatial and temporal variation of their food resources in these man-modified habitats, so that their potential to interact with other members of the granivore system is greatly reduced by human activities.     

Diverse foraging opportunities drive the functional response of local and landscape-scale bear predation on Pacific salmon

The relationship between prey abundance and predation is often examined in single habitat units or populations, but predators may occupy landscapes with diverse habitats and foraging opportunities. The vulnerability of prey within populations may depend on habitat features that hinder predation, and increased density of conspecifics in both the immediate vicinity and the broader landscape. We evaluated the relative effects of physical habitat, local, and neighborhood prey density on predation by brown bears on sockeye salmon in a suite of 27 streams using hierarchical Bayesian functional response models. Stream depth and width were inversely related to the maximum proportion of salmon killed, but not the asymptotic limit on total number killed. Interannual variation in predation was density dependent; the number of salmon killed increased with fish density in each stream towards an asymptote. Seven streams in two geographical groups with ≥23 years of data in common were then analyzed for neighborhood density effects. In most (12 of 18) cases predation in a stream was reduced by increasing salmon abundance in neighboring streams. The uncertainty in the estimates for these neighborhood effects may have resulted from interactions between salmon abundance and habitat that influenced foraging by bears, and from bear behavior (e.g., competitive exclusion) and abundance. Taken together, the results indicated that predator–prey interactions depend on density at multiple spatial scales, and on habitat features of the surrounding landscape. Explicit consideration of this context dependency should lead to improved understanding of the ecological impacts of predation across ecosystems and taxa.     

Multiple stages of tree seedling recruitment are altered in tropical forests degraded by selective logging

Tropical forest degradation is a global environmental issue. In degraded forests, seedling recruitment of canopy trees is vital for forest regeneration and recovery. We investigated how selective logging, a pervasive driver of tropical forest degradation, impacts canopy tree seedling recruitment, focusing on an endemic dipterocarp Dryobalanops lanceolata in Sabah, Borneo. During a mast‐fruiting event in intensively logged and nearby unlogged forest, we examined four stages of the seedling recruitment process: seed production, seed predation, and negative density‐dependent germination and seedling survival. Our results suggest that each stage of the seedling recruitment process is altered in logged forest. The seed crop of D. lanceolata trees in logged forest was one‐third smaller than that produced by trees in unlogged forest. The functional role of vertebrates in seed predation increased in logged forest while that of non‐vertebrates declined. Seeds in logged forest were less likely to germinate than those in unlogged forest. Germination increased with local‐scale conspecific seed density in unlogged forest, but seedling survival tended to decline. However, both germination and seedling survival increased with local‐scale conspecific seed density in logged forest. Notably, seed crop size, germination, and seedling survival tended to increase for larger trees in both unlogged and logged forests, suggesting that sustainable timber extraction and silvicultural practices designed to minimize damage to the residual stand are important to prevent seedling recruitment failure. Overall, these impacts sustained by several aspects of seedling recruitment in a mast‐fruiting year suggest that intensive selective logging may affect long‐term population dynamics of D. lanceolata. It is necessary to establish if other dipterocarp species, many of which are threatened by the timber trade, are similarly affected in tropical forests degraded by intensive selective logging.     

Density‐dependent pre‐dispersal seed predation and fruit set in a tropical tree

Negative density‐dependent demographic processes operating at post‐dispersal seed, seedling, and juvenile stages are the dominant explanation for the coexistence of high numbers of tree species in tropical forests. At adult stages, the effect of pollinators and pre‐dispersal fruit predators are often dependent on the density or abundance of flowers and fruit in the canopy, but each have opposite effects on individual realized reproduction. We studied the effect of density on total and mature fruit set and pre‐dispersal predation rates within individual tree canopies in a common canopy tree species, Jacaranda copaia in a 50‐ha forest census plot in central Panama. We sampled all reproductive sized trees in the plot (n = 188) across three years and estimated fruit set and predation rates. Population‐wide pre‐dispersal seed predation averaged between 6–37% across years. Using linear mixed effects models, we found that increased density and fecundity of conspecific neighbours increased focal tree fruit set, but also the rate of pre‐dispersal predation. An interaction between individual and neighbourhood fruit production predicted lower predation rates at high individual and neighbourhood fecundities, which suggests predator satiation at high fruit abundance levels. However, the rate at which fruit set increased with conspecific neighbour fruit production was greater than the rate at which fruit were lost to predation, resulting in an overall positive effect of neighbour density on mature fruit production in focal trees. Our results run counter to the expectation of a uniformly negative effect of density across all life stages in tropical trees and suggest further exploration of the role of spatial clumping, pollen dispersal limitation, and predation at pre‐dispersal adult stages in maintenance of species diversity in plant communities.     

Global Patterns of Pre-Dispersal Propagule Predation in Mangrove Forests1

Mangroves in disparate families produce viviparous seedlings (propagules) that are attacked by many crab and insect predators both before and after dispersal. While post-dispersal predation is viewed as an important factor in structuring many mangrove communities, pre-dispersal predation rates and agents have been characterized for few species. Ten species of mangrove and 3299 propagules were surveyed for pre-dispersal propagule predation at 42 sites around the world. Pre-dispersal predation rates were variable among sites and species, ranging from 0 to 93 percent within sands, with a global total predation rate of 23.3 percent (across all propagules examined) and a mean level of 28.3 percent across sites. Grapsid crabs, Coleoptera and Lepidoptera were the primary predators identified. Forests near human population centers and stands occurring at high intertidal sites exhibited higher levels of propagule predation than those in unpopulated or low-intertidal sites. Predation rates on a species were weakly, negatively correlated with conspecific seedling density at a site. To explore temporal variation in, and ramifications of pre-dispersal predation for propagule growth and abscission dynamics, Rhizophora mangle propagules were monitored over two years at three sites in Belize, Central America. Predation did not significantly reduce hypocotylar growth of germinated propagules on the parent tree, but nearly doubled the abscission rate of premature propagules. Pre-dispersal propagule predation is a ubiquitous feature of mangrove forests world-wide, and must be accounted for in estimates of reproductive output, stand health, and propagule availability for forestry and restoration efforts.     

Field observation of predation on adult lepidopterans by the tree sparrow Passer montanus saturatus

Many studies have examined defensive strategies of adult lepidopterans against bird predation. However, detailed field observations that describe how lepidopterans are hunted by wild birds are quite limited. I observed foraging behavior of wild tree sparrows on adult lepidopterans from May to June 2006 at Mt Yoshida, Kyoto, Japan. A total of 253 foraging attempts on at least 14 species of lepidopterans were observed. The tree sparrows attacked both resting and flying lepidopterans. When a tree sparrow approached a resting lepidopteran, the lepidopteran frequently flew to escape predation. The number of feeding attempts for resting and flying lepidopterans was 215 and 38, respectively. This result indicates that resting lepidopterans are the main target of predation by the tree sparrow. It is therefore considered that reducing the risk of detection is important for lepidopterans to avoid predation by the tree sparrow. Coloration as crypsis is probably effective in this respect.     

动物与植物种子更新的关系Ⅱ．动物对种子的捕食、扩散、贮藏及与幼苗建成的关系

植物的繁殖体总是面临来自各类生物（如昆虫、脊椎动物、真菌）的捕食风险。因动物捕食引起的种子死亡率影响植物的适合度、种群动态、群落结构和物种多样性的保持。种子被捕食的时间和强度成为植物生活史中发芽速度、地下种子库等特征的主要选择压力,而种子大小、生境类型等因素也影响动物对植物种子的捕食。捕食者饱和现象被认为是植物和种子捕食者之间的高度协同进化作用的结果,是限制动物破坏种子、提高被扩散种子存活率的一种选择压力。大部分群落中的大多数植物种子被动物扩散。种子扩散影响种子密度、种子被捕食率、病原体攻击率、种子与母树的距离、种子到达的生境类型以及建成的植株将与何种植物竞争,从而影响种子和幼苗的存活,最终影响母树及后代植物的适合度。种子被动物扩散后的分布一般遵循负指数分布曲线,大多数种子并没有扩散到离母树很远的地方。捕食风险、生境类型、植被盖度均影响动物对种子的扩散。植物结实的季节和果实损耗的过程也体现了其对扩散机会的适应。许多动物有贮藏植物种子的行为。动物贮藏植物繁殖体的行为,一方面调节食物的时空分布,提高了贮食动物在食物缺乏期的生存概率;另一方面也为种子萌发提供了适宜条件,促进了植物的扩散。于是,植物与贮食动物形成了一种协同进化关系,这种关系可能是自然界互惠关系（mutualism)的一种。影响幼苗存活和建成的因子包括种子贮蒇点的微生境、湿度、坡向、坡度、林冠盖度等。许多果食性动物吃掉果肉后,再将完好的种子反刍或排泄出来。种子经动物消化道处理后,发芽率常有所提高。     

Foraging patterns of voles at heterogeneous avian and uniform mustelid predation risk

Temporal variation of antipredatory behavior and a uniform distribution of predation risk over refuges and foraging sites may create foraging patterns different from those anticipated from risk in heterogenous habitats. We studied the temporal variation in foraging behavior of voles exposed to uniform mustelid predation risk and heterogeneous avian predation risk of different levels induced by vegetation types in eight outdoor enclosures (0.25 ha). We manipulated mustelid predation risk with weasel presence or absence and avian predation risk by reducing or providing local cover at experimental food patches. Foraging at food patches was monitored by collecting giving-up densities at artificial food patches, overall activity was automatically monitored, and mortality of voles was monitored by live-trapping and radiotracking. Voles depleted the food to lower levels in the sheltered patches than in the exposed ones. In enclosures with higher avian predation risk caused by lower vegetation height, trays were depleted to lower levels. Unexpectedly, voles foraged in more trays and depleted trays to lower levels in the presence of weasels than in the absence. Weasels match their prey's body size and locomotive abilities and therefore increase predation risk uniformly over both foraging sites and refuge sites that can both be entered by the predator. This reduces the costs of missing opportunities other than foraging. Voles changed their foraging strategy accordingly by specializing on the experimental food patches with predictable returns and probably reduced their foraging in the matrix of natural food source with unpredictable returns and high risk to encounter the weasel. Moreover, after 1 day of weasel presence, voles shifted their main foraging activities to avoid the diurnal weasel. This behavior facilitated bird predation, probably by nocturnal owls, and more voles were killed by birds than by weasels. Food patch use of voles in weasel enclosures increased with time. Voles had to balance the previously missed feeding opportunities by progressively concentrating on artificial food patches.