种子大小及其命运对植被更新贡献研究进展

种子是种子植物的繁殖体.种子对后代的资源投入将会直接影响后代的适合度进而影响到植被群落的更新.一般较大种子物种在种子萌发和幼苗生长阶段是具有较大优势的;但较小种子物种在逃避被动物采食和形成持久的土壤种子库成为植被更新的后备动力方面具有较大的优势.在不同的选择压力下,不同大小种子在以后的生活史中具有不同的命运,对植被的更新也具有不同的贡献.本文主要从种子大小和种子的传播散布、种子萌发、种子存活以及种子土壤库等方面综述了国外关于种子大小及其以后命运对植被更新贡献的研究结果,并对国内的相关研究提出了一些建议：种子生态学研究需要结合群落特点以及微环境等,开展从种子生产到幼苗更新全过程的系统性研究,为植物物种多样性保育和退化植被恢复方面提出有科学依据的管理措施.     

光照强度对六种草地植物种子萌发和幼苗生长的影响

光是影响种子萌发和幼苗生长的环境因素之一, 光照强度具有日动态和季节动态, 对植物出土和光合物质积累有重要影响。通过室内控制实验, 研究松嫩草地六种植物种子萌发及幼苗生长对光照强度的响应, 对深入理解松嫩草地植物生活史早期阶段的适应策略具有重要意义。研究结果表明: 防风(Saposhnikovia divaricate)、角蒿(Incarvillea sinensis)和黄金菊(Hypochaeris grandiflora)种子萌发率不受光照强度影响, 反枝苋(Amaranthus retroflexus)、大果琉璃草(Cynoglossum divaricatum)和紫菀(Aster tataricus)在弱光或中等光照下种子萌发率最高。六种种子的萌发指数在中等光强下最大, 且与无光处理间差异显著。除大果琉璃草和反枝苋在中等光强或弱光下胚根生物量最大外, 角蒿、防风、黄金菊、紫菀均在较强光下胚根生物量最大, 无光下最小; 大果琉璃草、黄金菊、防风、反枝苋的地上生物量不受光照强度影响, 而角蒿、紫菀在较强光下地上生物量最大; 角蒿和大果琉璃草在弱光或中等光强下根冠比最大, 防风、黄金菊、紫菀、反枝苋均在较强光下根冠比最大; 萌发率和萌发指数与种子大小之间没有显著相关性, 幼苗地上和胚根生物量与种子重量成显著正相关关系。光照强度对萌发率和种子大小之间的关系无显著影响, 但胚根生物量与种子大小之间关系的斜率受到光照强度影响, 中等光强下斜率最大。因此, 中等光照条件有利于松嫩草地植物种子萌发和幼苗建植。     

地中海沿岸沙丘种子大小对植物及其种子多度的影响

分析了地中海沿岸沙丘3种微生境(灌丛下、灌丛之间的开阔地、路径)以及整个沙丘生态系统的种子大小与土壤种子库中休眠种子的数量、土壤种子库中总种子的数量、地上植被中各个植物种的个体数量和幼苗数量、每种植物在小样方中的出现频率等方面之间的关系,也分析了种子尺寸(长、宽、高之和的平均值)与土壤种子的长久性、土壤种子库中休眠种子数量之间的关系;还分析了具有不同种子大小的植物在沙丘生态系统和各微生境的分布比率,验证了生活周期短的植物的种子在土壤中更长久和被干扰的微生境具有更多的休眠种子这种假说。结果发现,在地中海沿岸沙丘生态系统中,具有特别大的种子和特别小种子的植物种类都很少,不同植物种子的大小呈现近正态分布,但绝大多数植物的种子重量都不超过10 mg;微生境影响种子尺寸与种子长久性的关系,在灌丛下、开阔地区域等两个微生境以及整个的沙丘生态系统都呈现显著的正向关系(p<0 .0 5 ) ,而在路径这种关系不很明显;种子多度与植被物种出现频率呈现强烈的正相关(p<0 .0 5 ) ,没有发现种子大小与土壤种子库中休眠种子数量、种子库中总的种子数量、植被物种出现频率、植被的物种多度、幼苗植物多度等方面有显著的关系;具有中度大小种子(0 .1～10 mg)的植物在总的土壤种子库、休     

苔藓植物对植物天然更新的影响

种子是天然更新的基础,幼苗和幼树的生长是天然更新过程中最敏感的阶段。苔藓植物作为许多生态系统的主要地被物,在维管植物天然更新过程中发挥着不容忽视的作用。在许多植被类型中,都有发育良好的苔藓群落,大量的研究证明它们的存在影响到维管植物的萌发、建成和生长,并且可能进一步影响到物种的共生,有一些种类甚至可能指示树种的更新。本文就苔藓植物对物种更新早期阶段的作用和影响进行了综述。包括苔藓植物的生理特性造成的微气候变化;苔藓群落对种子传播、萌发及幼苗建成的影响。并讨论了苔藓植物与更新物种的种间关系及其对幼苗生长的化感作用。     

埋深和播种密度对豚草种子出苗及幼苗生长的影响

豚草是一种全球性的恶性入侵杂草,给我国生物多样性及经济发展造成了巨大威胁。为研究豚草种子出苗和幼苗生长的影响因素,采用盆栽实验分析了种子大小（L、M、S）与埋深（2、4 cm和6 cm）或播种密度（2、4粒/盆和8粒/盆）对豚草（Ambrosia artemisiifolia）种子的出苗和幼苗生长的影响。研究结果表明,埋深对豚草种子出苗的影响高于播种密度及种子大小的影响,较浅的埋深有利于豚草种子出苗;出苗率在不同播种密度下均具有较高水平,达到67.9%-100%,这种高出苗率是豚草在不同生境成功定植的原因之一。豚草的幼苗生长受埋深或播种密度的影响大于种子大小的影响,且小种子更易受埋深或播种密度的影响。种子大小显著影响豚草幼苗的株高和基径（P<0.05）,总体上较大种子的株高和基径高于小种子,此外更小的种子会将生物量更多的分配给根以促进幼苗的生长。幼苗的株高和基径在较浅的埋深下更高,而豚草幼苗的单株生物量、单株地上和地下生物量随埋深的增加而增加。播种密度的增加会加剧豚草幼苗之间对水分、营养、光照等资源的竞争,导致其单株地上、地下及总生物量显著降低（P<0.05）。研究发现豚草在浅埋深、低密度生境中更容易入侵成功,因此可以通过对土壤进行深翻,采取替代控制,种植竞争能力强的本地植物等手段有效管理和防治豚草。     

种子特征和播种深度对辽东栎种子萌发和幼苗生长的影响

该文在玻璃温室内的遮阴环境下,采用盆播方法研究了辽东栎种子特征(大小和种皮)和播种深度(0、3、6、10 cm)对种子萌发和幼苗生长的影响。结果表明:(1)辽东栎大种子萌发率、萌发速率、萌发值和萌发指数在所有播种深度均高于小种子。其中非去皮种子萌发率、萌发速率和萌发指数在0 cm播种深度的不同大小种子间差异显著,去皮种子所有萌发参数在6和10 cm播种深度的不同大小种子间均差异显著;去皮可促进大种子萌发,但抑制小种子萌发;不同大小种子所有萌发参数均在0 cm播种深度最大,在10cm播种深度最小。(2)不论有无种皮,大种子萌发幼苗的叶片数、单株叶面积、总干物质质量和根冠比在所有播种深度均大于小种子萌发幼苗;去皮种子萌发幼苗的株高、基径、叶片数、单株叶面积和总干物质质量在所有播种深度均不同程度地小于非去皮种子萌发幼苗,但前者根冠比在0、3、6 cm播种深度大于后者;随着播种深度增大,幼苗株高、叶片数、单株叶面积、总干物质质量和根冠比等生长参数均呈减小趋势,但基径随播种深度增大而增大。     

生物土壤结皮对种子萌发和幼苗建植的影响研究进展

生物土壤结皮（BSC）在陆地生态系统中具有重要的生态地位,尤其是旱地生态系统中,BSC占据了种子植物之间的广阔地面。因此,BSC的发展必然影响种子植物更新过程与植被空间格局;但其作用方式、影响程度等因相关研究涉及多气候要素、土壤类型、BSC组成物种和种子植物物种的差异及其不同组合,导致目前的研究结论存在广泛争议。研究综合论述了BSC改变地表微形态对种子传播过程的影响;BSC改变土壤特性（物理、化学、生物学）对种子萌发和幼苗存活与建植等关键环节的影响;并结合种子形态特征及种子萌发、幼苗建植的性状等,综合分析了BSC对种子传播、种子萌发与幼苗建植等更新过程的潜在影响机理;探讨了目前研究矛盾性结论产生的原因。总体来说,深入研究并全面揭示BSC对种子植物更新过程的影响,应加强学科交叉,将分子生物学、植物生理学、生物化学等微观研究,与遥感、野外生态因子过程监测、控制实验等宏观、中观研究结合,从机理到过程方面动态研究BSC对种子植物更新过程的影响,并引入水文模型、气候模型、种群动态模型等模型预测方法,研究气候变化、各类干扰频发的情景下,BSC对种子传播、萌发及幼苗建植过程的潜在影响,以期促进对BSC与种子植物间相互关系的研究,加深对干旱脆弱生态系统植被发展规律的认识。     

干扰强度对群落中紫茎泽兰种子萌发、幼苗定居和生长的影响

干扰与外来植物入侵密切相关,种子萌发和幼苗定居是植物生活史中最脆弱、也是外来植物入侵最关键的阶段.为研究干扰在恶性外来入侵植物紫茎泽兰成功入侵过程中的作用,采用人工牧草群落代替自然群落,人为干扰(去除不同面积的牧草)模拟自然干扰的方式,研究了紫茎泽兰入侵初期种子萌发、幼苗定居和生长对不同干扰强度的响应与适应.结果表明:...     

沙埋和种子大小对柠条锦鸡儿种子萌发、出苗和幼苗生长的影响

研究了沙埋深度和种子大小对内蒙古毛乌素沙地植被群落中占优势的柠条锦鸡儿(Caragana korshinskii Kom.)种子萌发、出苗、幼苗存活和生长的影响.结果表明,沙埋深度显著影响柠条锦鸡儿的种子萌发率、休眠率、出苗率、幼苗存活率及生物量.在0.5-2cm的浅层沙埋下,种子萌发率、出苗率、幼苗存活率及生物量最高,休眠率最低;沙埋深度≥4 cm时,柠条锦鸡儿的种子萌发率、出苗率、幼苗存活率及生物量随着沙埋深度增加显著降低,而休眠率却显著升高;沙埋深度≥12 cm时,柠条锦鸡儿种子不能够出苗,幼苗也不能够存活.种子大小对柠条锦鸡儿种子萌发率没有显著影响,但对出苗率、幼苗存活率及生物量影响显著.在各个沙埋深度下,不同大小的柠条锦鸡儿种子间的萌发率没有显著差异.当沙埋深度≤6 cm时,不同大小的柠条锦鸡儿种子在同一沙埋深度下的出苗率间没有显著差异;但当沙埋深度≥8 cm时,在同一沙埋深度下,大种子的出苗率显著高于中种子和小种子的出苗率,而中种子和小种子出苗率间没有显著差异.0.5-10 cm的沙埋深度中,除6 cm和8 cm深度下中种子和小种子萌发幼苗的生物量间没有显著差异外,其余深度下,大种子萌发的幼苗的存活率及生物量显著高于同一沙埋深度下中种子萌发的幼苗的存活率及生物量,后者又显著高于小种子萌发的幼苗的存活率及生物量.可能正是种子萌发对沙埋环境的忍耐或响应能力以及种子的多态性提高了柠条锦鸡儿在毛乌素沙地的适合度,为其在流动或半流动沙丘环境中成功定居并形成优势群落奠定了基础.     

广东亚热带森林木本植物幼苗生长特性研究

研究了广东亚热带42种木本植物幼苗的生长及其与物种的生态特性、生活型、种子大小的相互关系.较强光下(H,66.8 μmol m-2s-1)乔木幼苗的茎高和茎生物量显著高于灌木幼苗的相应值,但在较低光下(L,33.7 μmol m-2s-1)两者无显著差异.而阳性植物、耐阴植物和中间型植物之间,茎高和茎生物量无显著差别.乔木幼苗的叶面积和叶生物量比灌木幼苗大,但灌木幼苗的叶片数较乔木幼苗多.大种子种和小种子种幼苗之间,阳性植物、耐阴植物和中间型植物幼苗之间的叶片数和叶面积一般无显著差异.阳性植物幼苗比耐阴植物幼苗侧根数多.乔木幼苗的根生物量和根/茎比显著高于灌木幼苗在较高光下,阳性植物幼苗的根/茎比较耐阴植物幼苗高,但在较低光下无明显差异.45 d幼苗的根生物量与种子重量呈显著的正相关,而90 d幼苗无明显的相关.乔木幼苗个体生物量显著高于灌木幼苗.幼苗相对生长率和叶面积比的大小呈现如下顺序阳性植物＞中间型植物＞耐阴植物,但只有阳性植物和耐阴植物之间有显著差异.阳性植物、中间型植物和耐阴植物幼苗之间的单位叶率无显著差异.     

A comparative study of seed number, seed size, seedling size and recruitment in grassland plants

In this study we analyse relationships between seed number, seed size, seedling size and recruitment success in grassland plants. The often hypothesised trade-off between seed size and seed number was supported by a cross-species analysis and by an analysis of 35 phylogenetically independent contrasts, derived from a data-set of 72 species. Apart from among-species relatedness, we also controlled for possible confounding effect of plant size that may influence both seed size and seed number. A sowing experiment with 50 species was performed in the field. The seeds were sown in a grassland and subjected to two treatments, disturbance and undisturbed sward. Evidence for seed-limited recruitment was obtained for 45 of the species. Disturbance had a significant, or nearly significant, positive effect on recruitment for 16 of the 45 species. The relative recruitment in undisturbed sward increased with increased seed size, and both recruitment success and seedling size were positively related to seed size. We suggest that a trade-off between competitive ability and number of recruitment opportunities follows from the trade-off between seed size and seed number, through a causal chain from seed size via seedling size to recruitment success. The relationships between seed size, seed number and recruitment may be an important underlying mechanism for abundance and dynamics of plant species in grassland vegetation. This is an example of a direct link between evolutionary life-history theory, and theory of plant community structure.     

Effects of intraspecific and interspecific density on the demography of a perennial herb, Sanicula europaea

We examined the effects of intraspecific and interspecific competition on demographic processes in the perennial herb Sanicula europaea by manipulating the density of neighbouring plants. We followed the response in terms of survival, growth and reproduction and in terms of seedling recruitment. The demographic data from all phases of the life cycle enabled us to assess also the overall effects of treatments on population growth rate (λ) by transition matrix models. We also decomposed the differences in λ between control and treatments, using life table response experiments (LTRE). To study the effects of competition on recruitment in more detail and to evaluate the role of seed availability, we sowed seeds at different densities with or without vegetation removal.
Vegetative growth and flowering frequency of established individuals was not significantly affected by removal treatments, which suggest no, or a delayed response to released competition. Neighbour removal had no effect on seedling emergence but enhanced recruitment through a higher seedling survival. Conspecific and simultaneous conspecific and heterospecific removal of plants led to an increase in population growth rate (λ), whereas heterospecific removal alone led to a decrease. Emergence of seedlings and fate of vegetative established individuals contributed most to differences in λ between the control and the different treatments. Seed addition enhanced seedling emergence but, as seedling and juvenile survival were density dependent, densities of established individuals appear not to be seed limited.
In S. europaea removal treatments had different effects on established individuals and recruitment. This suggests that studies quantifying the effects of competition over the entire life cycle and performed in a natural environment are necessary to assess the importance of competition in perennial plant populations.     

Seedling recruitment in semi-natural pastures: the effects of disturbance, seed size, phenology and seed bank

We examined recruitment patterns in semi-natural pastures and their relationships to disturbance and species composition at different spatial scales (0.01 m² up to 4 m²) of both the vegetation and the seed bank. Possible associations between seedling recruitment and phenology were also studied. The study was performed in four pastures with different management history. Seedling recruitment was generally enhanced by disturbance, with the greatest effect for small-seeded species that germinate in the autumn. The local species-pool contributed to a large extent to seedling recruitment; 83 % of the recruited species were found in the vegetation, whereas 44 % of the recruited species were found in the seed bank. There were a total of 88 species found in the vegetation at different spatial scales; 61 % were recorded at the smallest spatial scale (0.01 m²). The area sampled at this scale comprised 1.25% of the area examined, indicating a small scale structuring of diversity in the pastures. The species number in the vegetation varied between 7 and 14 for the smallest scale (0.01 m²), and between 23 and 42 for the largest scale (4 m²), in the four pastures. The species number in the seed bank was less variable. Like most studies of perennial grasslands, we found no close correspondence between species distribution in the vegetation and in the seed bank. Two of the pastures had a joint management history and exhibited a close similarity of species in the seed bank, despite that one of the pastures was abandoned already in the beginning of this century.     

Positive relationship between fruit removal by animals and seedling recruitment in a tropical forest

Fleshy-fruited plants rely on animal frugivores to disperse their seeds, and seed removal by frugivores may leave an imprint on seedling recruitment. However, to what extent plant–frugivore interactions are related to seedling recruitment has rarely been quantified at the community level, especially in species-rich tropical forests. In this study, we tested the effect of different plant traits on fruit removal by frugivores and tested the relative importance of fruit removal, plant traits and abiotic factors for seedling recruitment. We quantified plant–frugivore interactions of 22 fleshy-fruited plant species consumed by 56 diurnal frugivore species, and counted the number of seedlings that emerged along an elevational gradient in the Colombian Andes. We measured a set of plant traits (i.e., crop size; fruit size; seed load and mass; fruit nutritional contents), estimated the density of adult plants and recorded relevant abiotic factors (light, temperature and humidity). We found that fruit removal by frugivores was positively associated with crop size, but negatively associated with fruit length and unrelated to seed load and fruit nutritional content. Seedling densities were positively related to the density of adult plants, seed mass and fruit removal by animals. We found no relationship between abiotic factors and seedling recruitment. Our results indicate that fruit abundance and morphology are important determinants of fruit removal and that fruit removal is positively associated with seedling recruitment accounting for effects of species abundance and plant traits. We conclude that plant traits shape fruit removal and seedling recruitment at the community level, while these two crucial processes of forest regeneration are directly linked by seed dispersal of animals.     

Interaction between large herbivore activities,vegetation structure,and flooding affects tree seedling emergence

Tree establishment in grazed vegetation mosaics involves a series of early bottlenecks, including seed dispersal, germination, seedling emergence, survival and growth. In a field experiment, we studied seedling emergence of two species with contrasting recruitment strategies, Fraxinus excelsior and Quercus robur, in five structurally different vegetations: grazed and ungrazed grassland, ruderal pioneer vegetation, soft rush tussocks, tall sedge mats and bramble scrub. In a simulation experiment, we studied the interaction effects of pre-emergence flooding (3 weeks of inundation), trampling and grazing (simulated by clipping) of grassland vegetation on the emergence and early growth of both tree species in grass swards. Seedling emergence was enhanced in low swards and sparse vegetation types. Despite different recruitment strategies, the interaction of flooding and trampling of swards enhanced seedling emergence of both species. Grazing of soft rush and tall sedges enhanced emergence of F. excelsior. Clipping grass swards increased early growth of emerging Q. robur. Our results support the hypothesis that natural disturbances of soil and vegetation create microsites for seedling emergence and reduce above-ground competition. In grazed systems however, these results suggest a discordant relationship between successful seedling emergence and subsequent seedling growth/survival during the establishment process in structurally different vegetations.     

Habitat degradation,topography and rainfall variability interact to determine seed distribution and recruitment in a sand dune grassland

Question: Understanding the mechanisms underlying how habitat degradation, topography and rainfall variability interactively affect seed distribution and seedling recruitment is crucial for explaining plant community patterns and dynamics. Interactions between these major factors were studied together in a semiarid sand dune grassland. Location: Eastern Inner Mongolia, China. Methods: The study system used four sites of fixed, semifixed, semishifting and shifting sand dune grasslands, representing a gradient of habitat degradation. We investigated the density of germinable seeds deposited in the top 5 cm of soil and in situ seedling emergence (number of seedlings emerging early in the growing season) and establishment (number of plants recruited at the end of the growing season) at three topographic positions (dune top, windward and leeward sides) within each site over 2 years that differed in rainfall. Habitat characteristics (i.e. vegetation cover, plant species composition and diversity, soil moisture and nutrient availability and soil erodibility) of the four sites were also measured. Results: Habitat degradation (i.e. decreased vegetation cover and enhanced wind erosion rate) significantly reduced the size of the germinable soil seed bank. On average, germinable seed number from the high‐vegetation cover fixed dune was 36‐fold larger than the low‐vegetation cover shifting dune, and eight‐ and two‐fold larger, respectively, than the semishifting and semifixed dunes with intermediate vegetation cover. We observed within‐habitat variability in seed distribution, but among‐topographic position variation differed among habitats. Seedling recruitment showed large between‐year, and among‐ and within‐habitat variability, but these variations varied significantly depending on the response variables evaluated (i.e. initial seedling density, final plant density, emergence rate and recruitment rate). Path analysis revealed complex density‐dependent positive and negative, direct and indirect effects of germinable seed density and initial seedling density on recruitment, but the relative importance of these density‐dependent effects varied depending on habitat type and rainfall availability. Conclusion: Our results suggest that habitat degradation, microtopography and rainfall availability interact in shaping sand dune seed bank and plant community recruitment patterns and dynamics. Their effects were mainly mediated through changes in both the biotic and abiotic environment during the process of habitat deterioration.     

Strong microsite control of seedling recruitment in tundra

The inclusion of environmental variation in studies of recruitment is a prerequisite for realistic predictions of the responses of vegetation to a changing environment. We investigated how seedling recruitment is affected by seed availability and microsite quality along a steep environmental gradient in dry tundra. A survey of natural seed rain and seedling density in vegetation was combined with observations of the establishment of 14 species after sowing into intact or disturbed vegetation. Although seed rain density was closely correlated with natural seedling establishment, the experimental seed addition showed that the microsite environment was even more important. For all species, seedling emergence peaked at the productive end of the gradient, irrespective of the adult niches realized. Disturbance promoted recruitment at all positions along the environmental gradient, not just at high productivity. Early seedling emergence constituted the main temporal bottleneck in recruitment for all species. Surprisingly, winter mortality was highest at what appeared to be the most benign end of the gradient. The results highlight that seedling recruitment patterns are largely determined by the earliest stages in seedling emergence, which again are closely linked to microsite quality. A fuller understanding of microsite effects on recruitment with implications for plant community assembly and vegetation change is provided.     

Effects of seed burial and fire on seedling and sapling recruitment, survival and growth of African savanna woody plant species

Seed predation and seedling mortality can act as strong demographic “bottlenecks” to sapling recruitment in African savanna woodlands. Fire also limits tree recruitment from saplings by suppressing their growth. I conducted field experiments with 13 woody plant species to assess the effects of seed burial on seedling emergence rates and effects of fire on seedling and sapling survival and growth rates over a period of 8 years at a savanna plot in central Zambia, southern Africa. Seed removal rates by small rodents varied among years and buried seeds had significantly higher emergence rates than seeds exposed to predators in most but not all the species. Annual burning reduced sapling growth in some species but in other species saplings experienced successive shoot die back even in the absence of fire. The findings show that for some woody species, seed predation is an important constraint to seedling recruitment but not for others and annual fires are important hindrances to demography and growth for some species but not others. Thus, demographic “bottlenecks” occur at different life history stages in different savanna woody species and these have the potential to alter woody tree competitive relationships and ultimately savanna structure.     

The Interplay among Acorn Abundance and Rodent Behavior Drives the Spatial Pattern of Seedling Recruitment in Mature Mediterranean Oak Forests

The patterns of seedling recruitment in animal-dispersed plants result from the interactions among environmental and behavioral variables. However, we know little on the contribution and combined effect of both kinds of variables. We designed a field study to assess the interplay between environment (vegetation structure, seed abundance, rodent abundance) and behavior (seed dispersal and predation by rodents, and rooting by wild boars), and their contribution to the spatial patterns of seedling recruitment in a Mediterranean mixed-oak forest. In a spatially explicit design, we monitored intensively all environmental and behavioral variables in fixed points at a small spatial scale from autumn to spring, as well as seedling emergence and survival. Our results revealed that the spatial patterns of seedling emergence were strongly related to acorn availability on the ground, but not by a facilitationeffect of vegetation cover. Rodents changed seed shadows generated by mother trees by dispersing most seeds from shrubby to open areas, but the spatial patterns of acorn dispersal/predation had no direct effect on recruitment. By contrast, rodents had a strong impact on recruitment as pilferers of cached seeds. Rooting by wild boars also reduced recruitment by reducing seed abundance, but also by changing rodent’s behavior towards higher consumption of acorns in situ. Hence, seed abundance and the foraging behavior of scatter-hoarding rodents and wild boars are driving the spatial patterns of seedling recruitment in this mature oak forest, rather than vegetation features. The contribution of vegetation to seedling recruitment (e.g. facilitation by shrubs) may be context dependent, having a little role in closed forests, or being overridden by directed seed dispersal from shrubby to open areas. We warn about the need of using broad approaches that consider the combined action of environment and behavior to improve our knowledge on the dynamics of natural regeneration in forests.