Scatterhoarding of Manchurian walnut <Emphasis Type="Italic">Juglans mandshurica</Emphasis> by small mammals: response to seed familiarity and seed size

Dispersal patterns can be affected by seed familiarity and seed traits, including size, mass, and nutritional value, but these factors have not been intensively studied in the context of seed dispersal processes. Our aim was to study how small rodents respond to seed size and seed familiarity in their pattern of Manchurian walnut (Juglans mandshurica) seeds in two different habitats in temperate forests of northeast China. Our results demonstrated that Apodemus penisulae acts as the most important disperser for Manchurian walnut seeds. Inexperienced small rodents did not reject seeds of the Manchurian walnut and show similar seed removal rates as compared with experienced rodents. Both experienced and naïve rodents actively participated in seed scatterhoarding of Manchurian walnut seeds. Consecutive survey showed that seeds with large size/mass were removed faster than those with small size/mass, indicating a preference for large seeds. However, small seeds scatter-hoarded by small rodents were transported farther than large ones, failing to support the traditional optimization models for various tree species. Small seeds of Manchurian walnut in caches were less likely to be recovered than large ones and showed greater cache survival rates, indicating that small seeds would be more advantageous for regeneration than large seeds in small rodent-dominated forests.     

Rare seed-predating mammals determine seed fate of <Emphasis Type="Italic">Canarium euphyllum</Emphasis>, a large-seeded tree species in a moist evergreen forest,Thailand

Natural seed deposition patterns and their effects on post-dispersal seed fate are critical to tropical tree recruitment. The major dispersal agents of the large-seeded tree Canarium euphyllum in Khao Yai National Park, Thailand, are large frugivorous birds such as hornbills, which generated spatially heterogeneous seed deposition patterns because they regurgitated seeds at perching trees and conspecific and heterospecific feeding trees. We investigated the fate of seeds dispersed in this manner using seed removal experiments and automatic camera trapping. Seeds placed experimentally around conspecific feeding trees had higher removal rates than seeds placed elsewhere. These effects were likely mediated by two seed-eating rodents, the Indochinese ground squirrel (Menetes berdmorei) and the giant long-tailed rat (Leopoldamys sabanus). Consequently, the spatial patterns generated by hornbills had consequences for post-dispersal seed fates, particularly whether or not the seeds were removed by rodents. Primary dispersal by hornbills does alter seed fate by altering the probability of rodent–seed interaction, but the ultimate impact of dispersal by hornbills will depend on how important rodent scatterhoarding is to seed germination and seedlings. Given that major seed dispersers of C. euphyllum are now absent or rare in degraded forests in tropical Asia, it is becoming increasingly important to understand the roles of scatterhoarding rodents in these altered habitats in this region.     

The Interplay of Habitat and Seed Type on Scatterhoarding Behavior in a Fragmented Afromontane Forest Landscape

Scatterhoarding by rodents, whereby seeds are collected and stored for later consumption, can result in seed dispersal. Seeds may be covered in litter on the forest floor (cached) or buried. This is particularly so in the Neotropics for large, nutritious seeds, and where primary dispersers are rare or missing. In African forests, contemporary anthropogenic pressures such as hunting, forest degradation, and fragmentation are contributing toward major declines in large frugivores, yet the potential for scatterhoarding to mitigate this loss is largely unknown. In this study, we used thread‐marked seed to explore the balance between seed predation and dispersal by rodents in Afromontane forest. We studied two tree species in three habitats: (1) continuous forest; (2) continuous forest edge, and (3) small, degraded riparian forest patches. We found that seed removal rates were high and almost the same in all three habitats for both tree species, but that the predation/dispersal balance differed among habitats. In continuous forest, more seeds of each species were scatterhoarded than depredated, and rates of scatterhoarding differed between the two species. In all habitats, burying seeds up to 2 cm belowground was more common than caching. Distances seeds were moved was approximately five times greater in continuous forest than in forest edge or riparian patches. We found strong evidence to suggest that the African pouched rat, Cricetomys sp. nov was responsible for the scatterhoarding.     

Survival and Scatterhoarding of Frugivores-Dispersed Seeds as a Function of Forest Disturbance

The effect of forest disturbance on survival and secondary dispersal of an artificial seed shadow (N= 800) was studied at Brownsberg Natural Park, Suriname, South America. We scattered single seeds of the frugivore‐dispersed tree Virola kwatae (Myristicaceae), simulating loose dispersal by frugivores, in undisturbed and disturbed secondary forest habitats. Seed survival rate aboveground was high (69%) within 2 wk and was negatively correlated with scatterhoarding rate by rodents, the latter being significantly lower in the undisturbed forest (9%) than in the disturbed forest (20%). Postdispersal seed predation by vertebrates was low (3%) and infestation of seeds by invertebrates was almost zero in all instances. Therefore, secondary seed dispersal by rodents in forest is not as critical for recruitment as observed among other bruchid‐infested large‐seeded species. Secondary seed dispersal by rodents may, however, facilitate seedling recruitment whether cached seeds experience greater survival than seeds remaining above ground surface.     

On the density-dependence of seed predation in<Emphasis Type="Italic"> Dipteryx micrantha</Emphasis>, a bat-dispersed rain forest tree

We studied the effect of seed density on seed predation by following the fate of bat-dispersed Dipteryx micrantha (Leguminosae) seeds deposited under bat feeding roosts. The study was conducted in Cocha Cashu biological station, Amazonian Peru, during the fruiting period of Dipteryx. Predation of Dipteryx seeds in the area is mainly by large to medium-sized rodents. Seed deposits beneath bat feeding roosts were monitored for a 13-week period in an 18-ha study area. A total of 210 seed deposits were found, and on average, seed predators encountered 22% of them during any one week. About one-third of the seed deposits escaped predation, and those deposits that had relatively few seeds were more likely to go unnoticed by rodents than were deposits with many seeds. The mean seed destruction rate was 8% per week; deposits with many seeds tended to lose a smaller proportion of their seeds to seed predators than did deposits with few seeds. Regression tests for the weekly data showed that, at the beginning of the observation period, seed predation was not density-dependent. Later, when the total seed crop beneath roosts was high, the number of seeds predated per deposit was positively density-dependent, while the proportion of seeds predated was negatively density-dependent, indicating predator satiation. Seed deposits that had been visited by seed predators once had a higher probability of being revisited the week after, especially if they contained many seeds when first encountered. This indicates that the foraging behavior of rodents may be affected by their remembering the location of seed-rich patches.     

Telemetric tracking of scatterhoarding and seed fate in a Central African forest

In seed predation studies, removal of a seed is only the first step of a dynamic process that may result in dispersal rather than seed death. This process, termed seed fate, has received little attention in African forests, particularly in Central Africa. We experimentally assessed the initial steps of seed fate for two tree species—the large‐seeded Pentaclethra macrophylla and the relatively small‐seeded Gambeya lacourtiana—in northeastern Gabon. Specifically, we evaluated whether seed size and seed consumer identity are important determinants of seed fate. We established experimental stations under conspecific fruiting trees, each comprising three seeds fitted with telemetric thread tags to facilitate their recovery, and a motion‐sensitive camera to identify visiting mammals. In total, animals removed 76 tagged seeds from experimental stations. Small Murid rats and mice primarily removed small Gambeya seeds, whereas large‐bodied rodents and mandrills primarily removed large Pentaclethra seeds. Gambeya seeds were carried shorter distances than Pentaclethra seeds and were less likely to be cached. The two large‐bodied rodents handled seeds differently: Cricetomys emini larderhoarded nearly all (N = 15 of 16) encountered Pentaclethra seeds deep in burrows, while Atherurus africanus cached all (N = 5 of 5) encountered Pentaclethra seeds singly under 1–3 cm of leaf litter and soil, at an average distance of 24.2 m and a maximum distance of 46.3 m from experimental stations. This study supports the hypothesis that seed fate varies based on seed size and seed consumer identity, and represents the first telemetric experimental evidence of larderhoarding and scatterhoarding in the region.     

Hoarding patterns of sigmodontine rodent species in the Central Monte Desert (Argentina)

Hoarding food is an important strategy of rodents in desert environments characterized by unpredictable and poor food resource availability. In the Monte Desert, Prosopis produces abundant food, unevenly in time and space, in the form of pods and seeds. Sigmodontine rodents (Graomys griseoflavus, Akodon molinae, Eligmodontia typus and Calomys musculinus) use Prosopis propagules extensively, and they could be predators or dispersers depending on how they handle and where they leave the propagules. The objectives of this study were: (1) to know what rodent species transported propagules; (2) to evaluate what hoarding pattern was used by species that transport propagules (larder and scatterhoarding); and (3) to analyse in which condition were propagules left by the rodent species, both at the food source and in caches. Our results showed that all four species transported propagules, with G. griseoflavus and E. typus being the species that carried more seeds. Our study supported the evidence that food caching is common among species and that many species both larderhoard and scatterhoard food. Graomys griseoflavus and A. molinae, the largest species, larderhoarded more than did the smaller E. typus. These results uphold the hypothesis that larger species will show greater propensity to larderhoard than smaller species. Considering the interaction between seed‐hoarding patterns and plants, E. typus was the species that could most improve germination because it scatterhoarded propagules and left seeds out of pods. In contrast, G. griseoflavus could have a negative impact on plant populations because this was the species that predated more seeds and larderhoarded a high percentage of them. The smallest C. musculinus was the species that transported propagules least, and left them as seeds inside pods or pod segments mainly at the food source, which makes seeds more vulnerable to predation.     

种子特征和结实量对啮齿动物取食和扩散种子行为的影响

为了深入了解啮齿动物在不同种子丰富度条件下对不同大小和单宁含量种子的觅食行为策略及其与植物种群更新的关系,在宁夏六盘山区的华北落叶松人工林,研究了不同大小和单宁含量[0%Tannin(T)、2%T、8%T和15%T]的人工种子在模拟结实小年和结实大年对啮齿动物取食和扩散行为的影响.结果表明: 啮齿动物消耗种子速度在结实小年更快,结实大年的种子消耗速度相对缓慢. 种子就地取食率(ISPR)在不同结实年份间无显著差异,扩散后取食率(PRAD)在结实小年显著高于结实大年,但前者的扩散后贮藏率(HRAD)显著低于后者;种子扩散后的取食距离(PDAD)和贮藏距离(HDAD)在结实小年均显著大于结实大年.在结实小年,大种子的PDAD和HDAD均大于小种子,前者在不同大小种子间均差异显著,而后者仅在2%T和15%T的不同大小种子间差异显著;在结实大年,除0%T外的其他单宁含量种子的PDAD和HDAD在不同大小种子间均差异显著.ISPR在中等单宁含量种子最大,高单宁含量种子最小;PRAD分别在结实小年的高单宁含量种子和结实大年的无单宁种子最大;不论在结实大年还是结实小年,HRAD均在高单宁含量种子最大,中等单宁含量种子最小.这说明结实大年可延缓啮齿动物对种子的消耗速率,提高种子的HRAD,但种子扩散距离减小;啮齿动物在结实大年和小年均表现出对大种子的扩散偏好,且大种子被扩散的距离更远;啮齿动物在不同结实年份均偏好于就地取食中等单宁含量种子,而扩散高单宁含量种子.     

Comparative recruitment patterns of two non-pioneer canopy tree species in French Guiana

Summary A comparative study was conducted on the recruitment patterns of two non-pioneer tree species, one dispersed by arboreal mammals and birds (Virola michelii, Myristicaceae) and the other by rodents (Moronobea coccinea, Clusiaceae). These species differ in fruiting phenology, seed size, dispersal distance, germination time and seed nutrient exhaustion. In both species, establishment patterns were consistent with the escape hypothesis and the Janzen-Connell model. Virola seeds need not be buried to survive and germinate, and may produce a seedling carpet beneath the parent. Moronobea seedlings only establish from seeds buried by scatterhoarding rodents in the surrounding understory. One-year survival of Virola seedlings was 47.8% and was greater >10 m than < 10 m from the largest parent tree. In contrast, survival of Moronobea seedlings was 56% 3 years after seed dispersal. Survival of juveniles was greater in gaps than in the understory for Virola but not for Moronobea. Moronobea survival was greater than Virola survival in both microhabitats. Both species establish in the understory, yet both grew faster in gaps. Virola appeared to be more gap-dependent than Moronobea which may persist several years in the understory until a gap occurs. Virola and Moronobea illustrate two intermediate recruitment patterns along an hypothetical continuum of nonpioneer species replacement (Bazzaz and Pickett 1980; Swaine and Whitmore 1988).     

Functional extinction of a desert rodent: implications for seed fate and vegetation dynamics

Population declines of once‐abundant species have often preceded understanding of their roles within ecosystems. Consequently, important drivers of environmental change may remain undiagnosed because we simply do not know how species that are now rare or extinct shaped ecosystems in the past. Australia's desert rodents are thought to have little numerical impact on seed fate and vegetation recruitment when compared with ants or with desert rodents on other continents. However most research on granivory by Australian desert rodents has occurred in areas where rodents were rare or functionally extinct. Here we ask if the paradigm that rodents are relatively un‐important granivores in Australian deserts is an artefact of their historical decline. In the Strzelecki Desert, the endangered rodent, Notomys fuscus is rare where introduced mesopredators are abundant but common where dingoes (an apex predator) suppress mesopredator populations. We used foraging trays to compare rates of seed removal for a common shrub (Dodonaea viscosa angustissima hopbush) between areas where N. fuscus, hopbush shrubs and their seedlings were rare and common and found that seed removal was consistently higher where rodents were common and hopbush rare. By excluding ants and rodents from foraging trays we show that ants removed more seeds than rodents where rodents were rare but rodents removed far more seeds than ants where rodents were common. By manipulating rodents’ access to the soil seed‐bank we show that hopbush seeds persisted in greater numbers where rodents were excluded than where they had access. Our results support the hypothesis that granivory by rodents may once have been a far more important process influencing the fate of seeds and shaping plant communities in arid Australia and suggest that dingo extirpation has cascading effects on shrub seeds. Our study highlights that functional extinction of rodents may be an under‐appreciated driver of vegetation change.     

鼠类对辽东栎橡子的搬运

在北京东灵山地区,在灌丛、森林2 种生境类型中共设计了5 个处理,以检验种子质量、种子可视度、生境类型如何影响鼠类对辽东栎（Quercus liaotungensis）橡子的搬运。每个处理包括200 枚种子。每日检查种子状态,实验持续16 d。鼠类在几天之内将大多数种子搬运走。完好橡子比虫蛀橡子消失得快。橡子在森林生境中明显比在灌丛生境中消失的快。鼠类对放置在灌丛下方和灌丛之间的橡子的搬运速率差别不大。放置在落叶下方的橡子消失速率明显慢于地表的橡子。本研究利用半存活时间测量种子的搬运率。各处理的半存活时间范围是0. 6 ～ 8. 6 d。各处理的橡子搬运率均较高,这说明鼠类有能力在种子下落到冬季来临的2 个月时间内,搬运绝大多数的橡子。实验说明,鼠类是辽东栎橡子重要的捕食者和扩散者。     

Spatial patterns of food storage by Merriam's kangaroo rats

We investigated scatter-hoarding behavior by Merriam's kangaroorats (Dipodomys merriami) in a complex arena in the laboratory.Animals cached seeds in four sand-filled boxes connected byrunways, with an artificial burrow at one end of the systemand a food source at the other. Animals initially cached moreseeds in boxes close to the food source, but amounts cachedbecame more evenly distributed among boxes as sessions progressed.The results were consistent with the hypothesis that scatterhoarding by kangaroo rats may represent a compromise betweenrapidly sequestering seeds, hence making them unavailable tonondigging competitors, and spacing-out caches, making themless vulnerable to pilferage by other rodents.     

Effect of plant cover on seed removal by rodents in the Monte Desert (Mendoza,Argentina)

Abstract In desert areas, predation risk is one of the highest costs of foraging and is a major influence on animal behaviour. Several strategies are used by foragers for surviving and reproducing in desert areas. The foraging strategies of the small mammals of South American deserts are still poorly known. In this study, we investigated the foraging strategies of rodents of the Monte Desert in response to distance from seed sources to sheltered sites (i.e. shrubs) during two different seasons (wet and dry). We evaluated the relative rates of removal of two species of seeds (millet and sunflower) by rodents at two sites by establishing 80 seed sources, 40 in unsheltered microhabitats and 40 in sheltered microhabitats. We recorded both the number of caches and seed consumption for each source. We found that plant cover affected the foraging activity of rodents of the sand dunes in the Monte Desert because both consumption and numbers of caches constructed from sheltered seed sources were higher than those from unsheltered ones. Consumption of sunflower was higher in the wet season than it was in the dry season, when millet consumption increased. Sunflower was the preferred seed both from sheltered or unsheltered sources. We discuss the possible causes of the different foraging strategies used by rodents of the Monte Desert.     

Directed vertebrate-mediated seed dispersal of a fleshy-fruited amaryllid in a heterogenous habitat

Most plants with fleshy fruits have seeds that are ingested by animals, but a less well-understood mode of seed dispersal involves fleshy fruits containing seeds that are discarded by frugivorous animals because they are too large or toxic to be ingested. We studied the seed dispersal biology of Haemanthus deformis, an amaryllid lily species found in a mosaic of bush clumps in a grassland matrix in South Africa. We asked whether seed dispersal is directed in and among bush clumps and whether germination and survival are greater for seeds dispersed to bush clumps than for those dispersed into grassland. Using camera trapping, we found that fruits are consumed mainly by birds and rodents. The pulp was removed from the seeds which were then discarded without ingestion. While many seeds were dispersed close to the parent plant, most (c. 78.5%) were dispersed further than 1 m away from the parent plant. Longer distance dispersal resulted mainly from birds flying off with fruits in their bill or from rodents engaging in scatter-hoarding behavior. Seedling survival was most successful within bush clumps as compared to grasslands and shade was identified as a primary requirement for seedling survival. Seeds from which the fruit pulp had been removed germinated faster than those in intact fruits. Haemanthus deformis deploys a system of directed seed dispersal, whereby both birds and rodents contribute to the dispersal of seeds within patchy bush clumps that are favorable for seedling survival.     

Granivory rates by rodents, insects, and birds at different microsites in the Patagonian steppe

We studied the rates of seed removal by different granivores, in different microsites, at different times, in a Patagonian shrub steppe in South America. Granivory rates of exotic ( Phalaris canariensis ) and native ( Mulinum spinosum ) seed species were an order of magnitude lower than those reported for another cool desert from North America with comparable climatic conditions. Insects and rodents removed the greatest amount of seeds in comparison to birds. In general, there were no differences in seed removal across microsites, except at one sampling time (when rodents and insects removed more seeds from bare soil and beside tussocks in comparison to positions beside shrubs, while birds took similar amounts of seeds from all microsites). The length of the experiment differentially affected the granivory rate of different groups. Removal rates (per day) were significantly greater, and exhibited lower variability, when seeds were left for a longer period of time in the field (a month) than for a few days. Insects were more efficient at finding the seeds rapidly and rodents at depleting them; birds could not find or deplete many seeds in short periods of time. Rates of granivory decreased slightly but significantly as the summer progressed mainly due to a reduction of seed removal by birds and rodents but not by insects. Granivores removed an order of magnitude less native seeds than exotic seeds. These differences seemed to be related to palatability as M. spinosum seeds have more phenols, toxic concentrations of iron and copper, and lower dry matter digestibility, phosphorous, and nitrogen content, in comparison to P. canariensis seeds.     

Seed preferences by rodents in the agri‐environment and implications for biological weed control

Post‐dispersal seed predation and endozoochorous seed dispersal are two antagonistic processes in relation to plant recruitment, but rely on similar preconditions such as feeding behavior of seed consumers and seed traits. In agricultural landscapes, rodents are considered important seed predators, thereby potentially providing regulating ecosystem services in terms of biological weed control. However, their potential to disperse seeds endozoochorously is largely unknown. We exposed seeds of arable plant species with different seed traits (seed weight, nutrient content) and different Red List status in an experimental rye field and assessed seed removal by rodents. In a complementary laboratory experiment, consumption rates, feeding preferences, and potential endozoochory by two vole species (Microtus arvalis and Myodes glareolus) were tested. Seed consumption by rodents after 24 h was 35% in the field and 90% in the laboratory. Both vole species preferred nutrient‐rich over nutrient‐poor seeds and M. glareolus further preferred light over heavy seeds and seeds of common over those of endangered plants. Endozoochory by voles could be neglected for all tested plant species as no seeds germinated, and only few intact seeds could be retrieved from feces. Synthesis and applications. Our results suggest that voles can provide regulating services in agricultural landscapes by depleting the seed shadow of weeds, rather than facilitating plant recruitment by endozoochory. In the laboratory, endangered arable plants were less preferred by voles than noxious weeds, and thus, our results provide implications for seed choice in restoration approaches. However, other factors such as seed and predator densities need to be taken into account to reliably predict the impact of rodents on the seed fate of arable 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.     

Seed dispersal of Korean pine Pinus koraiensis labeled by two different tags in a northern temperate forest, northeast China

Two types of tagging methods, i.e., a 1 × 3-cm tin tag attached to seed with a 10- to 12-cm metal wire (total weight 0.32 g) and a 2 × 4-cm white plastic tag fastened to seed with an identical metal wire (total weight 0.57 g) were used to study their effects on seed dispersal of Korean pine by small rodents. A total of 600 seeds were released to assess four main points: (1) difference in seed survival rates, (2) difference in caching behaviors of small rodents, (3) difference in dispersal distances, and (4) proportion of seed missing. The results demonstrated that seed removal for wire-plastic-tagged seeds was faster than that for wire-tin-tagged seeds. There was no apparent difference in the proportion of seeds eaten in situ (42% and 52% for wire-plastic-tagged seeds and wire-tin-tagged seeds, respectively). We found 41% and 1% of seeds were moved and hoarded for wire-plastic-tagged seeds and wire-tin-tagged seeds, respectively. However, 2.33% and 14% of seeds were missing, and their ultimate fates were not known for wire-plastic-tagged seeds and wire-tin-tagged seeds, respectively. We found the wire-plastic-tagged seeds easier to track than the wire-tin-tagged seeds due to the fact that the white plastic tags were more salient than the tin tags in field environments. The average dispersal distances were 4.11 ± 2.40 m and 3.01 ± 2.06 m for wire-plastic-tagged seeds and wire-tin-tagged seeds, respectively, and showed great difference. Despite most being eaten in situ or after removal, 41% of seeds were cached for wire-plastic-tagged seeds, much more than for wire-tin-tagged seeds. A total of 71 primary caches (123 seeds) were found for wire-plastic-tagged seeds, with the average and maximum cache sizes being 1.73 and 6, respectively. However, only three caches were found, and cache size was equal to one for wire-tin-tagged seeds. The above data suggests there is some uncertainty in different tagging methods to used track seed fates. Despite their effectiveness in helping to trace seed dispersal or movement by seed-dispersing rodents, different tagging methods—including size, color, and mass—need to be fully understand in enclosure experiments .     

Spatial patterns of acorn dispersal by rodents: do acorn crop size and ungulate presence matter?

Seed dispersal is qualitatively effective when it increases recruitment probability. A poorly studied factor likely affecting recruitment is the spatial distribution of dispersed seeds. Seed-caching animals are thought to disperse seeds in a way that reduces clumping and density to impede cache pilfering. Furthermore, dispersal might differ depending on whether the seed is immediately consumed versus cached for later consumption, and might differ depending on the ecological context. The main objectives of this study were to determine: 1) the spatial pattern of seed dispersal by rodents in a heterogeneous environment; 2) whether the patterns differ among years and among acorn competitor exclosure treatments, and 3) whether rodents create different spatial patterns of dispersal for acorns that are cached versus consumed immediately following dispersal. We studied the degree of spatial aggregation of acorn dispersal by rodents using two different estimators derived from the Ripley K and the Diggle G functions. We also analyzed various metrics of dispersal distances. For both analyses we used observed acorn dispersal patterns in two years differing in crop size and inside versus outside exclosures restricting access to acorn-consuming ungulates. During 2003, a year with a larger crop size, maximum seed dispersal distances were less, and the pattern of dispersed seeds was more clumped, than in 2004, a year with a smaller crop size. Median dispersal distances did not differ between years. In the presence of ungulates, seed dispersal was marginally sparser than in their absence. Cached acorns were dispersed more sparsely than acorns eaten immediately. These results have important implications for the quality of seed dispersal for oak recruitment that are likely relevant to other systems as well.     

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.