Re-caching of acorns by rodents: Cache management in eastern deciduous forests of North America

Scatter-hoarding rodents such as tree squirrels selectively cache seeds for subsequent use in widely-spaced caches placed below the ground surface. This behavior has important implications for seed dispersal, seedling establishment, and tree regeneration. Hoarders manage these caches by recovering and eating some seeds, and moving and re-caching others. This process of re-caching, however, is poorly understood. Here, we use radio-telemetry to evaluate re-caching behavior for the management of acorn caches by rodents in eastern deciduous forests. We also test the hypothesis that as seeds are re-cached, the distance from the source increases. Radio transmitters were implanted in Northern red oak (Quercus rubra) acorns and presented to rodents in a natural setting over 3 seasons. We used radio-telemetry to track and document evidence of recovery and re-caching. We tracked a total of 102 acorns. Of the 39 radio-tagged acorns initially cached, 19 (49%) were cached on two or more occasions; one acorn was cached four times. The hypothesis that rodents move seeds to progressively greater distances from the source is not well-supported, suggesting that acorns are being moved within an individual's home range. Given the species of rodents in the study area, gray squirrels (Sciurus carolinensis) are the most likely to be responsible for the caching and re-caching events. Gray squirrels appear to engage in extensive re-caching during periods of long-term food storage, which has important implications for understanding how caching behavior influences acorn dispersal and oak regeneration.     

Effectiveness of rodents as local seed dispersers of Holm oaks

In this study we assessed the effectiveness of rodents as dispersers of Quercus ilex in a patchy landscape in southeastern Spain. We experimentally followed the fates of 3,200 marked and weighed acorns from dispersal through the time of seedling emergence over three years. Rodents handled about 99% of acorns, and dispersed 67% and cached 7.4% of the dispersed acorns. Most caches were recovered and consumed, and only 1.3% of the original experimental acorns were found alive in caches the following spring. Dispersal distances were short (mean = 356.2 cm, median = 157 cm) and strongly right-skewed. Heavier acorns were dispersed further and were more likely to be cached and survive than lighter acorns. All caches were in litter or soil, and each contained a single acorn. Rodents moved acorns nonrandomly, mostly to oaks and pines. Most surviving acorns were either in oaks, a poor microhabitat for oak recruitment, or shrubs, a suitable microhabitat for oak recruitment. Our results suggest that rodents, by burying a relatively high proportion of acorns singly in shrubs and pines, act as moderately effective dispersers of Q. ilex. Nonetheless, this dispersal comes at a very heavy cost.     

Not only size matters: Acorn selection by the European jay (Garrulus glandarius)

A strong selection for acorn characteristics is expected to have evolved in the mutualistic relationship between the European jay (Garrulus glandarius) and the oak (Quercus spp.). Bossema's pioneer work suggested that jays do not select acorns randomly, but rather they preferentially select some size and species. Preference for some seeds over others may have implications on plant community dynamics by conferring advantages (or disadvantages) on the selected (avoided) seed characteristics. In this paper we test to what extent jays select acorns by species and/or by size and the relation between these two traits in Mediterranean oak species. The experiments consist of a set of field tests in which acorns from four different coexisting Mediterranean oak species (Quercus ilex, Quercus faginea, Quercus suber, and Quercus coccifera) were placed in artificial feeders accessible to wild jays. The acorns were previously measured to control individual acorn characteristics. Using video-recording techniques, we followed jay activity and the fate of each acorn (sequence of acorn selection and method of transport). Q. ilex acorns were preferred over other acorns, and Q. coccifera acorns were avoided when other acorns were available. Preference for Q. faginea and Q. suber acorns was intermediate, that is, they were preferred over Q. coccifera acorns but not over Q. ilex acorns. Large acorns were also preferred although acorn species selection was stronger than size selection. Jays selected species and size both by visual means and by using acorn area as an indicator of size. Acorns wider than 17–19 mm were carried in the bill because of throat limitation. Our results confirm Bossema's study on temperate oaks and extend it to Mediterranean oak species, revealing implications on mixed oak forest dynamics.     

Rodent acorn selection in a Mediterranean oak landscape

Quercus suber, Quercus ilex and Quercus coccifera (Cork, Holm and Kermes oaks, respectively) are common evergreen oak species that coexist in the landscapes of the western part of the Mediterranean basin. Rodents are the main acorn predators and thus one of the main factors for understanding recruitment patterns in oaks. In this paper we analyse to what extent mice prefer acorns from one oak species over another in three oak species studied using acorn removal experiments and video tape recordings. Twenty labelled acorns from each of the three Quercus species (60 acorns) were placed in 40 cm×40 cm quadrats on each plot. Because selection might vary as a result of the vegetation context, we performed the trials in the five main vegetation types within the study area (four replicates in each vegetation type) in order to control for habitat influences on rodent acorn preferences (a total of 20 plots). The removal of 1,200 acorns occurred within 68 days. Mice removed 98.7% of the acorns. Q. ilex acorns were preferred over Q. suber and Q. coccifera in all vegetation types except in pine forest, where no acorn preferences were detected. Acorn removal rates differed with vegetation type, correlating positively with shrub cover. The distance at which acorns were displaced by rodents (mean =4.6 m±5.1 SD) did not differ between acorn species, but varied among vegetation types. Bigger acorns of Q. coccifera were selected only after Q. ilex and Q. suber acorns were depleted, while no size selection was detected for the latter two species. Thus, we conclude that rodents show preference for some oak acorns and that landscape context contributes significantly to rodent activities and decisions.     

Seed predation and dispersal of glabrous filbert (Corylus Heterophylla) and pilose filbert (Corylus Mandshurica) by small mammals in a temperate forest, northeast China

We investigated the seed dispersal of glabrous filbert (Corylus heterophylla) and pilose filbert (Corylus mandshurica), two large-seeded shrub species in a temperate forest, northeast China, September 2006. Small mammals such as Apodemus speciosus, Clethrlonomys rufocanus, and Eutamias sibiricus, were regarded as the main dispersal agents. More seeds were harvested by small mammals in pilose filbert (98%) than in glabrous filbert (87.5%) till our last survey. Seed removal rates differed between the two species. Fewer seeds of glabrous filbert (17.5%) were eaten in situ than pilose filbert (57.5%). More seeds of glabrous filbert were removed (70%), stay intact after removal (25.5%), eaten after removal (16%) than pilose filbert. However, more seeds were cached after removal in pilose filbert than in glabrous filbert (10.5 and 4%, respectively). Fewer tagged seeds of pilose filberts (14%) were missed than glabrous filberts (24.5%). About 8 and 12 primary caches were found in glabrous filbert and pilose filbert seeds respectively, indicating scatter hoarding. All of the removed seeds were distributed within 10 m of seed stations for both filberts. The average dispersal distances for glabrous filbert did not differ from pilose filbert. Only a small proportion of the caches remained till our last survey (2 and 1%, respectively). Based on the results, we found a difference in dispersal patterns of glabrous filbert and pilose filbert seeds. Evidences showed that glabrous filberts might be a less preferred seed species for small seed-eating mammals compared with pilose filbert, probably due to its harder and thicker husk and low seed profitability.     

Guild structures of seed insects in relation to acorn development in two oak species

We investigated patterns of acorn growth in Quercus variabilis Blume and Quercus serrata Thunb., seasonal trends in emergence of seed insects found in the acorns, oviposition periods of the insects and falling periods of insect-infested acorns. In Q. variabilis, two insect guilds were associated with acorn development: (i) the immature acorn feeding (IAF) guild [Curculionidae sp., Poecilips cardamomi (Schaufuss), and Characoma ruficirra (Hampson)]; and (ii) the mature acorn feeding (MAF) guild [Curculio robustus (Roelofs), Curculio sikkimensis (Heller), and Cydia glandicolana (Danilevsky)]. In Q. serrata, there were three guilds: (i) the pistillate flower feeding (PFF) guild (cynipid wasp); (ii) the IAF guild [sap absorption by Mechoris ursulus (Roelofs)]; and (iii) the MAF guild (M. ursulus, C. sikkimensis, Cydia danilevskyi (Kuznetzov), C. glandicolana and Autostichidae sp.). The succession of guilds during acorn development may be a consequence of the use by different species of the limited food resource. The lack of a PFF guild in Q. variabilis that was found in our field site would have a positive effect on IAF guilds in utilizing the acorns.     

Scatter‐hoarding and cache pilfering of rodents in response to seed abundance

Seed caching and reciprocal cache pilferage play an important role in the coexistence of food‐hoarding animals. Understanding what affects seed caching and how cache pilferage occurs is an important question in seed dispersal ecology. However, tracking seed fate and cache pilferage presents substantial practical difficulties. Siberian chipmunks Tamias sibiricus always remove the entire pericarp when scatter‐hoarding acorns of Mongolian oak Quercus mongolica, whereas wood mice Apodemus peninsulae often store whole acorns in their caches. These differences in behavior provide an opportunity to investigate unilateral cache pilferage of T. sibiricus from A. peninsulae in response to seed abundance. In this study, tagged acorns were released at the peak and end periods of seed rain from Q. mongolica. This allowed us to investigate seed caching and unilateral cache pilferage at different seed abundances. We found that a higher proportion of acorns were cached at lower level of seed abundance (toward the end of seed rain), mainly because T. sibiricus rather than A. peninsulae scatter‐hoarded significantly more acorns at this time. Cache distances decreased with increasing seed abundance, indicating that acorns were cached further away and into smaller caches at lower seed abundance. Unexpectedly, unilateral cache pilferage by T. sibiricus was not significantly influenced by seed abundance—remaining at around 28% during both periods of high and low seed abundance.     

Hoarding behaviors of two wood mouse species: Different preference for acorns of two Fagaceae species

Two species of wood mouse, Apodemus argenteus and A. speciosus, were observed consuming and hoarding acorns of Quercus serrata and Castanopsis cuspidata. When each species of acorn was supplied individually, both species of mice used each species of acorn for eating and hoarding. When both species of acorn were supplied, A. argenteus consumed or hoarded only C. cuspidata, whereas A. speciosus tended to eat C. cuspidata acorns at the feeding site, and disperse or hoard Q. serrata acorns. Apodemus speciosus is unlikely to disperse C. cuspidata acorns (their utilization was biased towards consumption) when Q. serrata acorns are also available. Apodemus argenteus will make almost no contribution to the dispersal of Q. serrata when the two acorn species coexist.     

北京东灵山落叶阔叶林中辽东栎种子雨

在北京东灵山地区的一个落叶阔叶林中调查了辽东栎（Ｑｕｅｒｃｕｓ ｌｉａｏｔｕｎｇｅｎｓｉｓ Ｋｏｉｄｚ．）的种子雨。对于选定的４棵辽东栎中的３棵,树冠下的种子雨分布格局符合二次分布,具有很高的决定系数。由设置在树冠下的种子捕捉器收集的坚果数量来估计整棵树的种子雨。４棵树的种子雨中有活力的种子很少,变化范围从２６到２５９个。每棵树的树冠下的种子雨密度变化范围从０．７６到７．２６个／ｍ＾２。林中平均种     

Are tortoises important seed dispersers in Amazonian forests?

According to most studies on seed dispersal in tropical forests, mammals and birds are considered the main dispersal agents and the role played by other animal groups remains poorly explored. We investigate qualitative and quantitative components of the role played by the tortoise Chelonoidis denticulata in seed dispersal in southeastern Amazon, and the influence of seasonal variation in tortoise movement patterns on resulting seed shadows. Seed shadows produced by this tortoise were estimated by combining information on seed passage times through their digestive tract, which varied from 3 to 17 days, with a robust dataset on movements obtained from 18 adult C. denticulata monitored with radio transmitters and spoon-and-line tracking devices. A total of 4,206 seeds were found in 94 collected feces, belonging to 50 seed morphotypes of, at least, 25 plant genera. Very low rates of damage to the external structure of the ingested seeds were observed. Additionally, results of germination trials suggested that passage of seeds through C. denticulata’s digestive tract does not seem to negatively affect seed germination. The estimated seed shadows are likely to contribute significantly to the dispersal of seeds away from parent plants. During the dry season seeds were dispersed, on average, 174.1 m away from the location of fruit ingestion; during the rainy season, this mean dispersal distance increased to 276.7 m. Our results suggest that C. denticulata plays an important role in seed dispersal in Amazonian forests and highlight the influence of seasonal changes in movements on the resulting seed shadows.     

Satiation of predispersal seed predators: the importance of considering both plant and seed levels

Plants can reduce the fitness costs of granivory by satiating seed predators. The most common satiation mechanism is the production of large crops, which ensures that a proportion of the seeds survive predation. Nevertheless, satiation of small granivores at the seed level may also exist. Larger seeds would satiate more efficiently, enhancing the probability of seed survival after having been attacked. However, a larger seed size could compromise the efficiency of satiation by means of large crops if there were a negative relationship between seed size and the number of seeds produced by an individual plant. We analyze both types of satiation in the interaction between the holm oak Quercus ilex and the chestnut weevil Curculio elephas. Both crop size and acorn size differed strongly in a sample of 32 trees. Larger crop sizes satiated weevils, and higher proportions of the seeds were not attacked as crop size increased. Larger seeds also satiated weevil larvae, as a larger acorn size increased the likelihood of embryo survival. Seedling size was strongly related to acorn size and was reduced by weevil attack, but seedlings coming from large weeviled acorns were still larger. The number and the size of the acorns produced by individual trees were negatively related. Larger proportions of the crop were infested in oaks producing less numerous crops of larger acorns. However, contrary to expectations, these trees did not satiate more effectively at the seed level either. Effective satiation by larger acorns was precluded by larger multi-infestation rates associated to smaller seed crops, in such a way that the proportion of attacked seeds that survived did not vary among trees with different acorn sizes. These results highlight the need of considering satiation by means of large crops and large seeds in studies of predispersal seed predation. Long-term monitoring on individual oaks will help to assess whether there is a trade-off between the number and the size of the acorns and, if it existed, how it could condition the fitness consequences of both types of satiation.     

Spatial patterns in long-distance dispersal of Quercus ilex acorns by jays in a heterogeneous landscape

In this paper, I analyse the interaction between the holm-oak Quercus ilex , and one of its main dispersers, the European jay Garrulus glandarius , in an heterogeneous Mediterranean landscape. I quantify the spatial dispersal pattern of the seed shadow at two spatial scales, landscape (among patches) and microhabitat (within patches), by directly tracking the movement of seeds. Two main traits of the jay-mediated dispersal of holm-oak acorns across the landscape, the spatial pattern of dissemination and the distance from the source tree, are significantly and directly influenced by jay activity. Jays moved acorns nonrandomly, avoiding one main patch type of the study area to cache acorns, the shrubland-grasslands, and moving most of the acorns to pine stands, whether afforestation or open pinewoods. Within each patch type, jays had also a strong preference for caching acorns in some microhabitats, since>95% of the acorns dispersed by jays were cached beneath pines. The distance of holm-oak acorn dispersal was long in the study site, over 250 m, with some dispersals occurring up to 1 km from the source oaks. The shape of the dispersal kernel function fitted to the dispersal pattern produced by jays differed from those quantified for many other plant species. Jay-mediated dispersal had two components, one local and another produced by long-distance dispersal. Due to the heterogeneity of these Mediterranean environments, this difference in scale overlaps with a difference in habitat composition, short distances events resulting in dispersals within the same oak stands and long distance events resulting in dispersal outside of oak stands, usually to other vegetation units. Jay activity and movement pattern can have thus dramatic effects on both the local regeneration as well as the potential for regional spread of the holm-oak populations.     

Seed caching by woylies Bettongia penicillata can increase sandalwood Santalum spicatum regeneration in Western Australia

Abstract The role a small marsupial, the woylie Bettongia penicillata, might play in the recruitment and regeneration of Western Australian sandalwood Santalum spicatum through its seed caching behaviour was investigated in this study. To determine the fate of the seeds, cotton thread was attached to the seeds and the trail followed. A total of 25 seed caches were located. All of the seeds were found in separate caches, which was consistent with scatter‐hoarding behaviour. The average distance from the source of the seeds to the cache was 43.1 m ± 5.8 m at Dryandra woodland and 29.1 m ± 3.8 m at Karakamia sanctuary. The mean cache depth was 4.3 cm ± 0.2 cm at Dryandra woodland compared with 4.6 cm ± 0.3 cm at Karakamia sanctuary. Significantly more seedlings and saplings grew away from sandalwood trees at sites where woylies were present than at sites with no woylies. In contrast, significantly more seedlings and saplings grew under adult sandalwood trees at the site without woylies than where they were present, although there were significantly lower rates of recruitment and sandalwood regeneration at these sites. In addition, significantly more whole, undisturbed sandalwood seeds were found under the parent trees at the woylie‐free site than at the site with woylies. These findings strongly suggest that little seed dispersal or regeneration of sandalwood occurs in the absence of woylies. Through scatter‐hoarding, woylies have the potential to disperse and cache sandalwood seeds away from the source and significantly alter the subsequent regeneration of sandalwood. Furthermore, by caching seeds large distances away from a source, woylies could modify the distribution of sandalwood in an area.     

昆虫蛀蚀对鼠类介导下的锐齿槲栎种子扩散的影响

森林鼠类的种子贮藏行为对植物的扩散和自然更新有着非常重要的影响。然而,鼠类是否具有鉴别虫蛀种子的能力还存在一定的争议。此外,鼠类的鉴别能力是否受到食物丰富度变化的影响也未见相关报道。采用标签标记法,2011年秋季(9—11月,食物丰富季节)和2012年春季(4—6月,食物匮乏季节)分别在秦岭南坡的佛坪国家级自然保护区内,调查了森林鼠类对完好和虫蛀锐齿槲栎(Quercus aliena)种子的选择差异。结果显示:1)在秋季,尽管2种类型种子的存留动态没有显著差异,但是在后期虫蛀种子的存留时间相对更长;而在春季2种类型种子的存留动态则极为显著,几乎所有的完好种子(99%)在释放后的第3天就被鼠类全部扩散,虫蛀种子的存留时间则相对较长。2)在秋季,鼠类更喜好扩散后取食完好种子;而在春季,鼠类则喜好在原地取食绝大部分的种子,并且优先取食完好种子。3)在秋季,鼠类贮藏了更多的完好种子;而在春季,尽管完好种子在释放后第1天便达到贮藏高峰,然而由于后期的大量被捕食,2种类型种子在贮藏动态上没有显示出显著差异。研究结果表明秦岭地区森林鼠类可以准确区分完好与虫蛀种子,但是食物丰富度会影响鼠类对种子的选择策略。在食物丰富的秋季,鼠类更多地选择贮藏完好种子;而在食物相对匮乏的春季,鼠类更倾向于同时取食2种类型种子。森林鼠类通过对2种类型种子的鉴别和选择,影响不同种子的命运,从而可能对种子的扩散和自然更新产生重要影响。     

Rodents as seed dispersers in a heath — oak wood succession

Summary In a Danish heathland invasion of oak shrub has taken place, the succession rate being approximately 300 m during the last 100 years. The colonisation has occurred in steps related to the delay time between seedling stage and fertility stage. Seedlings are often found in clusters originating from caches probably made by seed-eating rodents. Apodemus sylvaticus, A. flavicollis and Clethrionomys glareoles. These rodents reached autumn densities of 25–50 individuals per ha. Radioactive acorns were scatterhoarded by the rodents, which mainly deposited the acorns singly up to at least 34 m from the oak shrub (mean 15.3±8.2 m), and preferably under Empetrum nigrum mats in the walls of runways Seedlings originating from radioactive acorns were found next summer at distances of 4–37 m from the oak shrub. In early summer caches containing new seedlings had a mean size of 2.0±2.2 acorns, range 1–16; mean distance of seedlings to nearest crown projection was 24.0±23.6 m, range 1–137. Thus, rodent acorn dispersal can explain the observed succession rate of oaks into the heathland     

Incorporating patterns of disperser behaviour into models of seed dispersal and its effects on estimated dispersal curves

The processes determining where seeds fall relative to their parent plant influence the spatial structure and dynamics of plant populations and communities. For animal dispersed species the factors influencing seed shadows are poorly understood. In this paper we test the hypothesis that the daily temporal distribution of disperser behaviours, for example, foraging and movement, influences dispersal outcomes, in particular the shape and scale of dispersal curves. To do this, we describe frugivory and the dispersal curves produced by the southern cassowary, Casuarius casuarius, the only large-bodied disperser in Australia’s rainforests. We found C. casuarius consumed fruits of 238 species and of all fleshy-fruit types. In feeding trials, seeds of 11 species were retained on average for 309 min (±256 SD). Sampling radio-telemetry data randomly, that is, assuming foraging occurs at random times during the day, gives an estimated average dispersal distance of 239 m (±207 SD) for seeds consumed by C. casuarius. Approximately 4% of seeds were dispersed further than 1,000 m. However, observation of wild birds indicated that foraging and movement occur more frequently early and late in the day. Seeds consumed early in the day were estimated to receive dispersal distances 1.4 times the ‘random’ average estimate, while afternoon consumed seeds received estimated mean dispersal distances of 0.46 times the ‘random’ estimate. Sampling movement data according to the daily distribution of C. casuarius foraging gives an estimated mean dispersal distance of 337 m (±194 SD). Most animals’ behaviour has a non-random temporal distribution. Consequently such effects should be common and need to be incorporated into seed shadow estimation. Our results point to dispersal curves being an emergent property of the plant–disperser interaction rather than being a property of a plant or species.     

Ficus seed shadows in a Bornean rain forest

Due to their copious seed production and numerous dispersers, rain forest fig trees have been assumed to produce extensive and dense seed shadows. To test this idea, patterns of seed dispersal of two species of large hemiepiphytic fig tree were measured in a Bornean rain forest. The sample included four Ficus stupenda and three F. subtecta trees with crop sizes ranging from 2,000 to 40,000 figs (400,000 to 13,000,000 seeds). Seed rain out to a distance of 60 m from each study tree was quantified using arrays of seed traps deployed in the understory. These trees showed a strongly leptokurtic pattern of dispersal, as expected, but all individuals had measurable seed rain at 60 m, ranging from 0.2 to 5.0 seeds/m². A regression of In-transformed seed rain density against distance gave a significant fit to all seven trees' dispersal patterns, indicating that the data could be fitted to the negative exponential distribution most commonly fitted to seed shadows. However, for six of seven trees, an improved fit was obtained for regressions in which distance was also In-transformed. This transformation corresponds to an inverse power distribution, indicating that for vertebrate-dispersed Ficus seeds, the tail of the seed rain distribution does not drop off as rapidly as in the exponential distribution typically associated with wind dispersed seed shadows. Over 50% of the seed crop was estimated to fall below each fig tree's crown. Up to 22% of the seed crop was dispersed beyond the crown edge, but within 60 m of the tree. Estimates of the maximum numbers of seeds which could have been transported beyond 60 m were 45% for the two largest crops of figs, but were under 24% for the trees with smaller crops. Seed traps positioned where they had an upper canopy layer above them were associated with higher probabilities of being hit by seeds, suggesting that vertebrate dispersal agents are likely to perch or travel through forest layers at the same level as the fig crown and could concentrate seeds in such areas to some degree. The probability of a safe site at 60 m from the fig tree being hit by seeds is calculated to be on the order of 0.01 per fruiting episode. Fig trees do not appear to saturate safe sites with seeds despite their large seed crops. If we in addition consider the rarity of quality establishment sites and post-dispersal factors reducing successful seedling establishment, hemiepiphytic fig trees appear to face severe obstacles to seedling recruitment.     

Rodents change acorn dispersal behaviour in response to ungulate presence

Small rodents are prominent seed predators, but they also favour plant recruitment as seed dispersers. The direct interactions of ungulates on plants are more one‐sided and negative, as they mainly reduce plant recruitment through predation on seeds and seedlings. The effects of small rodents and ungulates on plant recruitment have been considered and studied as independent episodes within plant regeneration cycles. However, ungulate–rodent interactions and their potential effects on plant regeneration have not been considered so far. A number of studies have recently documented ungulate effects on the abundance, diversity and spatial distribution of small rodents. Here, we hypothesize that ungulates may also affect rodent seed dispersal behaviour. We monitored acorn dispersal by small rodents (Mus spretus and Apodemus sylvaticus) in oak woodlands with and without exclosures for large ungulates, mainly red deer, Cervus elaphus, and wild boar, Sus scrofa. The study was carried out in a typical Mediterranean Holm oak, Quercus ilex, forest throughout the acorn fall season in 2003 and 2004. We found that, in both years, the proportion of acorns cached and not recovered in the short‐term was, on average, lower in the presence (1.4%) than in the absence (19.9%) of ungulates. Acorn dispersal distances were not affected by ungulate presence in either year. However, ungulates had an effect on the spatial distribution of dispersed seeds; rodents apparently avoided shrubs as caching sites in both years. This result was interpreted as a behavioural response to reduce the risk of cache pilferage by conspecifics, which are closely associated with shrubs in presence, but not in absence, of ungulates. Potential effects of different densities of rodents or predators were discarded, as none of them differed between the areas with and without ungulates. The present study found significant interactions between heterospecific seed and seedling consumers that had been considered as independent episodes within tree regeneration cycles. As a result of such interactions, ungulates may have negative indirect effects on oak recruitment by reducing (1) acorn caching frequency, and (2) the proportion of acorns cached under shrubs, key nurse‐plants for the establishment of Holm oak seedlings in Mediterranean areas.     

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.     

岩松鼠和松鸦对辽东栎坚果的捕食和传播

脊椎动物对栎属(Ｑｕｅｒｃｕｓ)坚果的捕食和传播是栎属通过实生苗更新的关键因子,栎属中一些物种坚果的分布格局以及动物对坚果的捕食作用已有较多研究[1～6]。一般情况下,栎属的坚果成熟落地后立即开始发芽,落在地表的坚果里的种子胚根长出后,由于地表枯落物的阻碍作用...