Alternative strategies of seed predator escape by early-germinating oaks in Asia and North America

Early germination of white oaks is widely viewed as an evolutionary strategy to escape rodent predation; yet, the mechanism by which this is accomplished is poorly understood. We report that chestnut oak Quercus montana (CO) and white oak Q. alba (WO) (from North America), and oriental cork oak Q. variabilis (OO) and Mongolian oak Q. mongolica (MO) (from Asia) can escape predation and successfully establish from only taproots. During germination in autumn, cotyledonary petioles of acorns of CO and WO elongate and push the plumule out of the cotyledons, whereas OO and MO extend only the hypocotyls and retain the plumule within the cotyledons. Experiments showed that the pruned taproots (>6 cm) of CO and WO acorns containing the plumule successfully germinated and survived, and the pruned taproots (≥12 cm) of OO and MO acorns without the plumule successfully regenerated along with the detached acorns, thus producing two seedlings. We argue that these two distinct regeneration morphologies reflect alternative strategies for escaping seed predation.     

The choice of seed tracking method influenced fate of beech seeds dispersed by rodents

We tracked seeds of European beech (Fagus sylvatica) dispersed by rodents in Gorzowska Forest (western Poland). We used two seed labeling methods, marking with UV-fluorescent powder and with plastic tags, to test whether using different marking methods influences results of seed tracking. The removal rates did not differ among seeds marked with UV-powder, seeds labeled with tags, and unmanipulated seeds. We found 78 % of removed seeds marked with tags, but only 25 % of UV-marked seeds. The consumption rates of tagged and UV-marked seeds were dramatically different: rodents ate 83 % of the former and 26 % of the later. The average dispersal distance was larger for seeds marked with UV-powder than for tagged seeds. Our findings suggest that the choice of seed tracking method might influence results of seed dispersal studies.     

Seed dispersal and predation in the endemic Atlantic rainforest palm <Emphasis Type="Italic">Astrocaryum aculeatissimum</Emphasis> across a gradient of seed disperser abundance

Tropical forests have been subject to intense hunting of medium and large frugivores that are important in dispersing large-seeded species. It has been hypothesized that in areas with extinction or low abundance of medium and large-bodied animals the density of small rodents may increase. Therefore, this increment in the density of small rodents may compensate for the absence or low abundance of medium and large frugivores on seed removal and seed dispersal. Here, we fill up this gap in the literature by determining if seed removal, seed dispersal, and seed predation by small rodents (spiny rats, Trinomys inheringi and squirrels, Sciurus ingrami) are maintained in defaunated areas. We accessed seed removal, seed dispersal, seed predation, and seedling recruitment of an endemic Atlantic rainforest palm, Astrocaryum aculeatissimum, in a gradient of abundance of agoutis. We found that seed removal, scatter hoarding, and seed predation increase with the abundance of agoutis. In contrast, the proportion of dispersed but non-cached seeds decreased with the abundance of agoutis. We did not find any effect of the abundance of agoutis on seed dispersal distance, but we did find a positive trend on the density of seedlings. We concluded that small rodents do not compensate the low abundance of agoutis on seed removal, scatter hoarding, and seed predation of this palm tree. Moreover, areas in which agoutis are already extinct did not present any seed removal or scatter hoarding, not even by small rodents. This study emphasizes both the importance of agoutis in dispersing seeds of A. aculeatissimum and the collapse in seed dispersal of this palm in areas where agoutis are already extinct.     

Growth and survival of cork oak (Quercus suber) seedlings after simulated partial cotyledon consumption under different soil nutrient contents

Aims

We examined the importance of partial seed consumption (cotyledon loss) by rabbits in the early establishment of seedlings of cork oaks restricted to nutrient-impoverished soils.

Methods

To determine the importance of cotyledons in the growth and development of seedlings, we simulated two levels of predation [light (30 % cotyledon loss) and heavy (60 % loss) partial consumption] and two soil nutrient contents (nutrient-poor soil, nutrient-rich soil). Seedlings height, root length, dry root and shoot biomass, specific leaf mass, leaf density, gas exchange, chlorophyll fluorescence parameters and photosynthetic pigment concentrations were determined.

Results

Results indicated that effect of nutrient level on the growth of the oak seedlings was more important than that of cotyledon biomass. However, in nutrient–poor soils, cotyledon biomass played a major role in the early performance of cork oaks. Acorns grown in nutrient-rich substrate, despite having greater aerial vigor, were slower to develop a vertical root, and hence less likely to reach permanent moisture. Cotyledon loss caused a decrease in the biomass of roots and shoots when acorns were heavily consumed, and as a result experienced a reduction in net photosynthetic rate, stomatal conductance and chlorophyll concentration. Survival of seedlings was unaffected by either soil type or cotyledon loss.

Conclusions

Our results show that effects of soil type on the survival of oak seedlings were more important than those of cotyledon biomass. However, in a competitive situation, cotyledon biomass, as an indicative of growth nutrient support rather than an energy source, could be vital in a nutrient-poor environment, particularly in Mediterranean climate regions and for species with little inherent drought tolerance (as is the case of Quercus spp.), where rapid root growth is required to ensure that contact with soil moisture is maintained over the first summer.     

Incorporating insect infestation into rodent seed dispersal: better if the larva is still inside

Many nutritious seeds are commonly attacked by insects which feed on the seed reserves. However, studies have not fully explored the ecological implications of insect infestation in animal seed dispersal and subsequent plant regeneration. Our question is whether the fact that an infested seed still contains the larva or not might increase/decrease the probability of being successfully dispersed by animals. This study examines the effects of weevil-infested seeds on the natural regeneration of a rodent-dispersed oak species. Rodents showed a high ability to discriminate between sound and infested seeds, even when the larva was still inside. As a result, rodents caused differential seed dispersal for sound and infested seeds by modifying multiple aspects of the dispersal process. We found that, for the same seed weight, infested acorns with a larva still inside can contribute to natural regeneration (0.7?% of seedlings in next summer), although in comparison to sound acorns they suffered higher predation rates by rodents (both partial and complete), were removed later from the ground (less preferred), cached less frequently, and dispersed to shorter distances, which reduced their potential to colonize new environments. However, infested seeds with exit holes are notably less preferred by rodents and, when dispersed, they are mostly deposited on the litter (uncached) with shorter dispersal distances and lower emergence success. Thus, the probability that larval-holed acorns will produce viable seedlings is extremely low (null in this study). Whether infested seeds still contain a larva or not clearly determines the probability of being successfully dispersed. Premature seed drop prolongs the presence of the larva inside the acorn after seed drop, and could be a possible mechanism to allow dispersal of infested seeds.     

啮齿动物对不同林木种子的搬运和取食微生境选择机制

不同植物种子依靠不同的方式实现扩散,啮齿动物对林木种子搬运后在取食点微生境和贮藏方式的选择存在偏好,研究其贮藏行为与微生境的关系是探究幼苗建成的关键。在秦岭中段火地塘林区,采用标签标记法,以锐齿槲栎、华山松和油松种子为材料,探究了小型啮齿动物对松栎混交林建群种种子扩散过程的影响。结果表明:1)油松种子原地取食率显著高于锐齿槲栎和华山松种子,且啮齿动物更倾向于搬运后取食(60%)和埋藏(4.33%)华山松种子,搬运后取食距离也为华山松最大(2.49 m);锐齿槲栎小种子被搬运后埋藏的距离最大(4.92 m)。2)除华山松种子外,其他类型种子被搬运后单个取食的比例均在85%以上;油松种子不存在埋藏点,而其他类型种子90%以上均以单个形式被埋藏。3)大部分种子被啮齿动物搬运后选择在裸地丢弃;锐齿槲栎大种子(87.5%)、小种子(78.57%)和华山松种子(53.33%)较大比例被啮齿动物埋藏在灌丛下方,埋藏在裸地的种子较少。4)大部分种子在灌丛下方被取食,仅华山松种子被啮齿动物搬运到洞穴取食;除油松种子被大量原地取食外,其他类型种子被搬运到取食点的种子比例基本呈现由微生境植被复杂到简单(灌丛—草丛—灌丛边缘—裸地)而逐渐减小的趋势。种子的营养价值及取食和搬运过程中啮齿动物付出的成本是影响种子命运的关键性因子,且啮齿动物对种子埋藏和取食地点的微生境存在较明显的选择性。     

The Proximate Basis of the Oak Dispersal Syndrome: Detection of Seed Dormancy by Rodents

Previously we have shown how a range of physical and chemicalcharacteristics of acorns influences the behavioral decisionsof food-hoarding rodents which in turn affects the dispersal,establishment and spatial arrangement of oaks. One such behaviorinvolves the selective caching of acorns of red oaks (subgenus:Erythrobalanus) over those of white oaks (Quercus) because ofreduced perishability that results from delayed germinationof acorns in the red oak group. In this study, we sought toidentify the specific proximate cues (visual and olfactory)that eastern gray squirrels (Sciurus carolinensis) use whenmaking these decisions. In two series of field experiments,we presented individual, free-ranging animals with pairs ofexperimentally altered acorns (that differed with respect toa single chemical or visual characteristic) and recorded theirfeeding and caching responses. Squirrels cached artificial acornswith pericarps (shells) of red oak acorns and ate those withshells of white oak regardless of the internal chemical compositionof either type of acorn. Only when the shells of artificialacorns were first soaked in acetone (to remove potential chemicalodors) did animals eat artificial acorns made with the shellsof red oak acorns. Squirrels also ate one-year old red oak acornsthat had broken dormancy, even when they exhibited no signsof germination. We argue that a chemical cue in the shell ofacorns is important in the detection of seed dormancy and thedecision to cache acorns, and that such a cue might ultimatelycontribute to the differential dispersal of red and white oaksby rodents.     

Postdispersal Seed Fate of Tropical Montane Trees in an Agricultural Landscape, Southern Costa Rica

Variation in postdispersal seed fate is an important factor driving patterns of forest regeneration. Because most previous studies have not tracked final seed fate and have commonly equated seed removal with predation without considering the possibility of secondary dispersal, little is known about individual seed mortality factors in successional and degraded habitats. This study tracked the postdispersal fate of large-seeded tropical montane trees in abandoned pasture, young secondary forests, and small forest fragments during two consecutive years in an agricultural landscape in southern Costa Rica. The incidence of secondary dispersal by animals, scatterhoarding in particular, and the effects of seed burial on germination were measured. Overall, seeds survived through germination more often in secondary forests with high levels of mortality occurring in abandoned pastures and forest fragments. The majority of seed mortality was caused by rodent predation in forest fragments, insects and fungal pathogens in secondary forests, and a combination of desiccation, insects, and fungal pathogens in pastures. Seeds were frequently secondarily dispersed in larger forest fragments, whereas they were only rarely moved in pastures and secondary forests. Burial tended to improve germination in pastures and was important for an often scatterhoarded species, Otoba novogranatensis, in all habitats. The results of this study suggest that: (1) seed mortality factors differ in response to the type of habitat degradation; (2) large-seeded species have high potential for survival when dispersed to young secondary forests; and (3) seed removal is not a reliable proxy for seed predation, particularly in less degraded forest fragments.     

六盘山区辽东栎的实生苗更新及其影响因子

研究了六盘山区辽东栎(Quercus liaotungensis)灌丛的种子产量、土壤种子库组成、湿沙和风干贮藏对种子寿命的影响以及动物取食子叶对种子萌发和幼苗建立的影响。结果表明, 辽东栎灌丛种子的完好率为27.51%, 被动物取食或搬运种子的比例(41.51%)显著高于其他类型种子(p < 0.01); 辽东栎次生林土壤种子库中萌发和虫蛀种子分别占35.16%和38.29%, 完好种子仅占13.65%, 捕食动物主要通过贮藏或搬运而影响土壤种子库中的种子密度。湿沙贮藏60天的辽东栎种子自动萌发率高达96.67%, 短期贮藏可加快种子的萌发进程, 提高萌发率和萌发指数, 但随着贮藏时间的延长, 种子萌发进程延迟, 萌发率、萌发指数和活力指数均不同程度地降低。排除动物取食处理的幼苗在林窗和林下生境的存活率分别为80%和83%, 而不排除动物取食幼苗在2种生境中分别仅有25%和31%能够存活, 表明子叶在幼苗建立中具有重要作用。林窗中幼苗子叶的动物取食率(85.00%)高于林下(71.00%), 子叶留存的幼苗在林窗中的存活率(6.00%)低于林下(15.50%), 而子叶被取食幼苗的存活率在两种环境中基本相等(分别为18.50%和18.00%)。     

Fragmentation modifies seed trait effects on scatter-hoarders’ foraging decisions

Scatter-hoarding animals are crucial in seed dispersal of nut-bearing plants. We used the holm oak Quercus ilex—wood mouse Apodemus sylvaticus mutualism as a model system to evaluate the relative importance of seed size and fat content on scatter-hoarders’ foraging decisions influencing oak dispersal and potential recruitment. We performed a field experiment in which we offered holm oak acorns with contrasting seed size (2 vs 5 g) and fat content (3 vs 11%). Moreover, to test if the strength of these seed trait effects was context-dependent, experimental acorns were placed in small fragments, where natural regeneration is scarce or absent, and forest habitats. In small fragments, rodents had to face increased intraspecific competition for acorns and reduced anti-predator cover during transportation. As a result, they became more selective to ensure rapid acquisition of most valuable food items but, in turn, transported seeds closer to avoid unaffordable predation risks. During harvesting and caching, larger acorns were prioritized and preferentially cached. Fat content only had a minor effect in harvesting preferences. In contrast, in forest sites, where rodent abundance was four times lower and understory cover was well-developed, rodents were not selective but provided enhanced dispersal services to oaks (caching rates were 75% higher). From the plants’ perspective, our results imply that the benefits of producing costly seeds are context-dependent. Seed traits modified harvesting and caching rates only when rodents were forced to forage more efficiently in response to increased intraspecific competition. However, when landscape traits limited cache protection strategies, a more selective foraging behavior by scatter-hoarders did not result in enhanced dispersal services. Overall, our result shows that successful dispersal of acorns depends on how specific traits modulate their value and how landscape properties affect rodents’ ability to safeguard them for later consumption.     

An Advantage of Large Seed Size: Tolerating Rather than Succumbing to Seed Predators1

Although large seeds might be more attractive and apparent to seed predators, large seed size could enable tolerance of seed predators. If seeds are large enough to sustain damage that would kill smaller seeds yet still produce viable seedlings, investment above the minimum by the maternal plant could be advantageous. I tested this hypothesis by removing 0–80 percent of the cotyledons of four large-seeded (4–180 g) tree species from Papua New Guinea and monitoring germination and seedling growth for eight months. All species showed little negative effect on seedling size with up to 50 percent removal of cotyledons and the larger species showed a less serious effect on growth than smaller-seeded species above 50 percent removal. Large-seeded species clearly have more than the minimum-required cotyledonary reserves. Observations of viable seedlings with heavily damaged cotyledons suggest that these species withstand attack by rodents and beetles by virtue of their large size.     

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.     

Impacts of secondary seed dispersal and herbivory on seedling survival in Aesculus turbinata

Abstract. Aesculus turbinata is a tree species with large seeds (6.2 g mean dry weight). We studied the demography of its seeds and seedlings in a temperate deciduous forest in northern Japan to elucidate the ecological significance of large seeds with special reference to herbivory and secondary dispersal. Both seed and seedling stages suffered greatly from herbivores. Seedling herbivory was important judged from experiments with shoot clipping and hypogeal cotyledon removal. However, some seedlings survived through re-sprouting after herbivory. Survival rate and percentage resprouting seedlings were lower than those with remaining cotyledons, though seedling size was not affected. This suggests that stored resources in hypogeal cotyledons are working as a kind of ‘risk hedge’ against severe aboveground shoot clipping experienced by A. turbinata. The spatial distribution of seedlings was expanded via seed scatter-hoarding by rodents. Seedling survival rate was higher within canopy gaps than under closed canopy, indicating that canopy gaps are safe sites for establishment, and was negatively correlated with seedling density. Therefore, secondary seed dispersal in this species seems to be effective in ‘finding’ safe sites and in ‘escaping’ density-dependent mortality. The large seeds and seedlings of A. turbinata are attractive to herbivores, but the high resistance of seedlings to herbivory due to large reserves and the effective secondary dispersal appear to mitigate these disadvantages.     

Acorn removal and dispersal by the dung beetle Thorectes lusitanicus: ecological implications

Abstract.  1. Plant–animal interactions, and in particular the processes of seed predation and dispersal, are crucial for tree regeneration and forest dynamics. A novel and striking case of interaction between a dung beetle ( Thorectes lusitanicus ) and two Quercus species ( Q. suber and Q. canariensis ) in forests of southern Spain is presented here.
2. During the autumn, T. lusitanicus beetles (endemic to the southern Iberian Peninsula) bury and feed on single-seeded fruits (acorns) of Quercus , with important ecological implications.
3. Field experiments found differences in the removal rate of acorns by T. lusitanicus , depending on the type of microsite within the forest, the species of oak, the exclosure of large herbivores, and the forest site.
4. Acorn consumption by T. lusitanicus was studied under laboratory conditions, confirming for the first time that this dung beetle is a legitimate seed predator.
5. In addition, some buried acorns can be abandoned partially predated or even intact, and emerge as seedlings; thus, T. lusitanicus also has a potential role as secondary seed disperser.     

Importance of microhabitat and acorn burial on Quercus ilex early recruitment: non-additive effects on multiple demographic processes

This study investigates the effect of microhabitat and seed burial on the main demographic processes operating during the early recruitment of Quercus ilex, such as postdispersal seed predation, seed germination, and seedling emergence, survival and growth. The effect of burial was positive over all the processes analysed in this study, since predation rate was lower (63.6% vs. 88%), whereas germination (53.1% vs. 21.8%) and emergence (32.0% vs. 5.5%) were higher for buried acorns. The quality of some microhabitats remained similar throughout the stages and processes studied. Thus, afforestation provided especially suitable microhabitats for oak establishment, since seed predation was lower, while germination, emergence and seedling survival were higher, than in any other microhabitat. By contrast, the quality of some microhabitats, such as open sites and Holm oaks, differed between recruitment stages. Acorns in open sites escaped predation and germinated easily, but most seedlings died due to summer drought. Similarly, although acorns under Holm oaks can germinate and survive drought, they cannot survive to postdispersal predators. This uncoupling results in a post-dispersal change in the spatial distribution of Q. ilex recruits. Furthermore, there were significant interactions between burial and microhabitat for some demographic processes. The recruitment was in afforestations high irrespective of burial, suggesting that burial is not as beneficial in high-quality habitats as it is in lower-quality ones. An accurate understanding of plant recruitment requires the determination not only of the direct effects of limiting factors but also the potential interactions occurring between them.     

生境对啮齿动物介导下锐齿槲栎种子命运的影响

干扰作为森林演替的驱动力,在改变森林结构和地表植被的同时也对林内动物介导下的种子扩散过程产生影响.本研究采用塑料标签标记法,选取秦岭林区不同人为干扰后形成的3种类型生境(未抚育林分、抚育3年后林分、裸地),研究啮齿动物扩散行为对不同生境类型锐齿槲栎种子命运的响应.结果表明:抚育3年后林分的种子扩散速率最大,裸地扩散速率最小.裸地的种子原地取食率显著低于未抚育(25.0%)和抚育3年后林分(36.3%),而抚育3年后林分的种子搬运后取食率显著高于未抚育林分(17.3%)和裸地(5.0%),且抚育3年后的林分中被啮齿动物分散埋藏的种子比例最高(4.3%).种子扩散距离以抚育3年后林分最大(26 m),且扩散距离显著大于裸地和未抚育林分.锐齿槲栎种子的初次扩散过程在不同生境中表现不同,其种子后期成苗率也存在差异.不同类型生境在一定程度上影响啮齿动物的觅食策略,最终导致不同的种子扩散模式和森林天然更新格局的形成.     

滇青冈和高山栲种子昆虫寄生及其对种群更新的影响

对昆明西山地区滇青冈(Cyclobalanopsis glaucoides)和高山栲(Castanopsis delavayi)种子(坚果)虫蛀率在两个年份、整个落种期和山坡不同部位的变化情况进行研究,结果表明：① 寄生于滇青冈种子内的象甲为青冈象(Curculio megadens)和Curculio sp.,寄生于高山栲种子内的象甲是Curculio sp.和剪枝象(Cyllorhynchites sp.);② 滇青冈样地的虫蛀率为2484%~2760%,样地间无显著差异,两年间的平均虫蛀率亦无显著差异;高山栲样地间虫蛀率为1794%~2655%,有显著差异性,两年间的平均虫蛀率也有显著差异。③ 高山栲的下落种子虫蛀率随时间逐步下降,9月和10月中下旬出现两个虫蛀率的高峰期;高山栲种子内寄生的昆虫落地后即入土化蛹,滇青冈落地种子内的寄生昆虫在初期会滞留一段时间;④ 虫蛀率与林下实生苗数量无显著相关性,实生苗多集中在大树周围以及草本层植物较丰富的小生境。     

Do seed predation and dispersal limit downslope movement of a semi-desert grassland/oak woodland transition?

Abstract. Semi-desert grasslands and dryland forests are typically arranged along elevation gradients, with low elevation grasslands and savannas separated from higher-elevation woodlands by a diffuse boundary. Recent (< 200 yr) woody plant encroachment into adjacent semi-desert grasslands appears to be a cosmopolitan phenomenon, and has been attributed to disturbance by humans (e.g., livestock grazing, fire suppression); however, little is known about rates, patterns and mechanisms. Are observations of increased woody plant abundance in semi-arid grasslands indicative of a downslope shift in the boundary zone? Experimental plots were established in adjacent oak woodlands and semi-desert grasslands in southeastern Arizona, to determine if mechanisms of acorn predation and dispersal would facilitate or limit downslope movement of oak woodlands. We recovered acorns that had been dispersed 50 m into adjacent grasslands, which suggests that they were cached by acorn predators. Acorns in grassland plots were more than twice as likely to escape predation than acorns that remained in woodland or cleared woodland plots. Contrary to previously published research, simulated perturbations (gap formation, understory and below-ground vegetation removal) did not affect seed predation, suggesting that generalizations from mesic, deciduous oak forests may not be accurately extrapolated to xeric, evergreen oak forests. Recently published studies indicate that xeric conditions limit survival of oak seedlings across the boundary zone, suggesting that observed patterns of acorn dispersal and predation provide the potential for colonization of adjacent grassland only during infrequent years with cool, moist conditions during late summer months.     

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     

Tree-shrub associations in grazed woodlands: first rodents,then cattle?

Spatial associations of tree saplings with spiny or toxic plants in grazed woodlands are generally explained by associational resistance, i.e., protection against grazing via a well-defended neighbor. In this study, we tested whether directed seed dispersal and post-dispersal seed removal by wood mice are additional explanations for the observed spatial association between thorny shrubs and trees, hence before associational resistance. We performed three studies in grazed woodlands in the Netherlands to test this idea. Our first seed dispersal experiment with tagged acorns showed indeed that wood mice disperse acorns directed towards shrubs. The majority of these dispersed acorns were, however, consumed. Our second experiment revealed that post-dispersal removal of cached acorns was higher under shrubs than in grassland and under trees, but also indicated the importance of within-shrub position: shrub centers were faster depleted than shrub edges. Also the number of freshly emerged seedlings and older saplings differed between micro-habitats, being higher under trees and at shrub inner-edges than at shrub outer-edges, shrub centers, and in grassland. The spatial associations with shrubs got stronger with the age of saplings, which probably reflect accumulated grazing effects over time, being higher in the open than in shrubs. We conclude that directed acorn dispersal and post-dispersal acorn removal by wood mice are two additional explaining mechanisms for the spatial associations between thorny shrubs and trees in grazed woodlands, before associational resistance. Our findings change the view that large herbivores are the sole main drivers behind tree recruitment patterns in grazed woodlands.