Large acorns benefit seedling recruitment by satiating weevil larvae in <Emphasis Type="Italic">Quercus aliena</Emphasis>

Plants can enhance their seed viability and survival through reducing the fitness costs of herbivory by satiating seed predators at either plant or seed level, or both. The former satiates with production of large crops while the latter with large-sized seeds, enhancing the probability of seed survival. We analyzed both types of satiation in the interaction between the Quercus aliena and the weevil Curculio spp in 2 years with different crop size and acorn size. Larger crop size seemed to satiate less effectively weevils due to higher proportions of the seeds were attacked in oaks producing more crops of smaller acorns. However, prematurely abscission of infested acorns in large crop year might compensate this ineffectiveness. Larger acorns in lean year well satiated weevil larvae, as a larger acorn size increased the likelihood of embryo survival and seedling establishment. Although effective satiation by larger acorns would be negated by smaller seed crops, the proportion of attacked acorns that survived increased with large acorn size in lean year. These results highlighted the importance of annual variation in acorn size of Quercus aliena in satiating and defending weevil larval predations, consequently in successful seedling recruitment and establishment.     

Factors limiting the recruitment of Quercus schottkyana,a dominant evergreen oak in SW China

Quercus schottkyana is a dominant species of oak in the Asian evergreen broad-leaved forests in southwestern China but seedlings are uncommon and recruitment is rare. Annual acorn production by Q. schottkyana is variable and the acorns are exposed to a series of mortality risks. Understanding the factors that limit recruitment of the oak requires knowledge of the oak's life cycle from acorn production to germination and seedling establishment. In this study, we first tested the effects of acorn density on establishment of seedlings by placing batches of acorns at different densities throughout the study area. Second, we tested the effects of herbivores on seedling survival by erecting fences around both natural and transplanted seedling populations. Our results show that even though the rate of seedling establishment increases as acorn density increases (for 32-8000 acorns·m^-2), survival rates of seedlings in the field were generally low (0-0.6%). We show that seedling recruitment of Q. schottkyana is mainly limited to the acorn stage where 88% of the acorns died from the combined effects of desiccation and predation by weevils (Curculio) and bark beetles (Coccotrypes sp.). Herbivory results in the death of some seedlings and consequently also affects the recruitment of seedlings of Q. schottkyana.     

Fruit abortion,developmental selection and developmental stability in<Emphasis Type="Italic"> Quercus ilex</Emphasis>

Fruit abortion has been hypothesized to be a parental means of selective removal of propagules with low viability. In particular, aborted zygotes have been suggested to have developmentally deviant phenotypes, and surviving offspring may therefore give rise to adults with a developmentally stable phenotype. We tested predictions from this hypothesis using acorns of holm oaks Quercus ilex as a model system. Fecundity of oak trees was negatively related to mean fluctuating asymmetry of leaves, and abortion rates were positively related to leaf fluctuating asymmetry in at least one population. Aborted acorns were asymmetric in 83-99% of cases in three samples, while mature acorns were only asymmetric in 57-78% of cases. Acorn asymmetry was unrelated to germination probability and germination date, and had no significant effect on number of leaves, leaf mass, stem mass, seedling height or leaf area of seedlings. However, acorn asymmetry affected the trade-off between number and size of leaves in seedlings. Seedlings from asymmetric acorns showed a positive relationship between acorn size and number of leaves, but no relationships between acorn size and leaf area, while symmetric acorns showed the opposite. A positive relationship between acorn size and number of leaves in spring was found for naturally emerged seedlings that died during their first summer, whereas the number of leaves produced by surviving seedlings did not depend on acorn size. These findings are consistent with the hypothesis that stressed trees selectively abort propagules of low viability, and that developmental selection acts on a measure of developmental instability of fruits.     

The impact of the common rabbit on cork oak regeneration in SW Spain

Seed predation has traditionally been considered as simply a progressive loss of seeds. However, partial consumption of tree seeds may actually assist regeneration. We quantified partial consumption of acorns by the common rabbit (Oryctolagus cuniculus) in a cork oak forest in southwest Spain (Do?ana Natural Park). To examine the effects of partial consumption on germination and seedling emergence, we simulated two different levels of acorn predation (30 and 50?% cotyledon loss, representing light and heavy partial consumption, respectively). One-third of the sampled acorns presented evidence of consumption but most of the damage caused by partial consumption did not affect the acorn embryo (damage was mainly to non-vital parts of the propagule). Partial consumption caused more rapid germination and emergence than that was observed in intact acorns, but germination and emergence rate was unaffected. Heavy consumption of the acorn caused differences in root and shoot biomass as a consequence of cotyledon loss. In this greenhouse study, partial consumption was non-lethal to the acorns, and cotyledon biomass loss did not decrease the probability of successful establishment. Partially consumed seeds can therefore contribute significantly to the natural regeneration of Quercus coccifera, although it would be beneficial to conduct long-term studies focusing on seed traits and animal behaviour under field conditions.     

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.     

Acorn crop size and pre-dispersal predation determine inter-specific differences in the recruitment of co-occurring oaks

The contribution of pre-dispersal seed predation to inter-specific differences in recruitment remains elusive. In species with no resistance mechanisms, differences in pre-dispersal predation may arise from differences in seed abundance (plant satiation) or in the ability of seeds to survive insect infestation (seed satiation). This study aimed to analyse the impact of pre-dispersal acorn predation by weevils in two co-occurring Mediterranean oaks (Quercus ilex and Quercus humilis) and to compare its relevance with other processes involved in recruitment. We monitored the patterns of acorn production and acorn infestation by weevils and we conducted experimental tests of acorn germination after weevil infestation, post-dispersal predation and seedling establishment in mixed forests. Monitoring and experimental data were integrated in a simulation model to test for the effects of pre-dispersal predation in recruitment. In both oaks pre-dispersal acorn infestation decreased with increasing acorn crop size (plant satiation). This benefited Q. ilex which exhibited stronger masting behaviour than Q. humilis, with almost a single and outstanding reproductive event in 6 years. Acorn infestation was more than twice as high in Q. humilis (47.0%) as in Q. ilex (20.0%) irrespective of the number of seeds produced by each species. Although germination of infested acorns (seed satiation) was higher in Q. humilis (60%) than in Q. ilex (21%), this could barely mitigate the higher infestation rate in the former species, to reduce seed loss. Conversely to pre-dispersal predation, no inter-specific differences were observed either in post-dispersal predation or seedling establishment. Our results indicate that pre-dispersal predation may contribute to differences in seed supply, and ultimately in recruitment, between co-existing oaks. Moreover, they suggest that seed satiation can barely offset differences in seed infestation rates. This serves as a warning against overemphasising seed satiation as a mechanism to overcome seed predation by insects. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

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.     

High Acorn Predation Prevents the Regeneration of Quercus liaotungensis in the Dongling Mountain Region of North China

The mature oak (Quercus liaotungensis Koidz) forests in the Dongling mountains of northern China have become degraded in recent years because regeneration has been limited. To determine whether or not seedling establishment of the oak is seed limited, microsite limited, or predator limited and to determine whether seedling establishment is affected by ground cover, we conducted field experiments during a mast year and investigated the fate of seeds and the soil seed bank dynamics of the oak. A large acorn crop (128.8 acorns/m²) was observed in the study period, and the peak density of acorns on the forest floor reached 46.5 acorns/m², suggesting that tree recruitment was not seed limited. Acorns in the soil seed bank were mainly lost through decay (principally after fungal attack), consumption in situ, and removal by animals. Predation (including consumption in situ and removal) accounted for 86.4% of acorn loss and was therefore likely to have been the most important factor influencing seed dynamics. More than 70% of acorns were found to have germinated, but no established seedling was observed on the forest floor. Using cages to exclude predators, it was estimated that 87% of acorns germinated and 49% became established as seedlings, indicating that the acorns on the forest floor could emerge and grow in the absence of predators. We conclude that the regeneration of the tree population is limited by predators rather than by the availability of microsites. The presence of ground cover increased the germination rate and increased the chance of seed survival in the early stage of the experiment, but at the end of the investigation, no established seedling was found in the quadrats both with and without ground cover, possibly because of high density of animal predators. On the basis of these results, we suggest that selective tree felling will increase the coverage of the herbaceous layer, which can further decrease the population density of the rodents, and thereby improve the regeneration of oak trees.     

Effect of Acorn Burying Depth on Germination,Seedling Emergence and Development of Quercus aliena var. acuteserrata

Acorns of Quercus aliena var. acuteserrata Maxim. are often predated by small mammals and birds in natural forests. These animals not only eat the acorns during the acorn ripening season, but also cache and hoard most of the remaining acorns on the forest floor in the soil for their future use. These buried acorns form the main seed resource for regeneration. Burying depth is potentially important for germination and for seedling development. The effects of burying depth on germination and seedling development in relation to acorn size were studied in an experiment, in which acorns were planted at 6 cm-, 12 cm- and 18 cm-depth. The experimental results showed that fewer acorns germinated as burying depth increased. From the deeply buried acorns fewer seedlings emerged at later time than from those acorns buried less deeply. They appeared to have more difficulties to emerge above-ground than the seedlings from shallowly buried acorns. The deeply buried acorns and their seedlings also appeared to be more susceptible to rot. Acorn size did not significantly affect germination and emergence of the seedlings. As early emerged seedlings had longer developmental periods in their first growing season, and therefore grew better than the late emerged seedlings, seedlings from the shallowly buried acorns took the advantage.     

锐齿槲栎橡子埋藏深度对发芽、幼苗出土和发育的影响

锐齿槲栎（Quercus aliena var.acuteserrata Maxim.)林内活动的许多型哺乳动物和鸟类不仅采食橡子,而且还常窖贮或在土壤大量的橡 子以便季后利用。这些埋藏的橡子构成了发育有效更新苗的主要种子来源。橡子被埋藏的深度和橡子的大小对幼苗的发育有潜在的影响。把橡子埋藏在6、12和18cm3个土层深度的实验结果显示：随着埋藏深度的增加,橡子的发芽率和发芽橡子的幼苗出土率都有明显下降,由深埋橡胶子发育的幼苗出土比较困难,所需时间较长,也更易腐烂,橡子的大小对发芽和幼苗出土没有明显的影响,早出土幼苗在第一个生长季内由于有较长的生长发育期,季末发育得较好,所以,由浅埋橡子发育的幼苗在更新成功方面有优势。     

东灵山地区动物对辽东栎坚果的捕食和传播Ⅰ.排除啮齿目动物对坚果丢失的影响

动物对辽东栎(Quercus liaotungensis Koidz.)坚果的捕食被认为是影响辽东栎坚果命运的主要因素,因此直接影响幼苗的建立和自然更新。在东灵山一个落叶阔叶林中调查了辽东栎坚果被脊椎动物转运和就地消耗状况。在排除部分小型啮齿目动物前后,辽东栎坚果的丢失动态曲线不同。到实验结束时,基本上所有放置的辽东栎坚果都消失了。排除动物前的曲线上有一个拐点,而排除动物后的丢失曲线上有多个拐点。两曲线相似的地方是：最初一两天辽东栎坚果的丢失非常迅速。排除小型啮齿目动物对辽东栎坚果的丢失有影响,然而坚果尺寸和微生境条件对坚果的丢失没有影响。任意2个因子的交互作用以及3个因子的交互作用对辽东栎坚果的丢失也都没有影响。排除啮齿目动物和坚果尺寸对辽东栎坚果的就地消耗有影响,微生境以及其它因子的交互作用对坚果的就地消耗没有影响。在9月份3d捕鼠的时间里,共捕到啮齿目动物3种：大林姬鼠(Apodemus speciosus)、社鼠(Niviveniter confucianus )和花鼠(Tantias sibiricus)。研究结果表明,尽管啮齿目动物的排除能够降低辽东栎坚果的丢失速度,但它们(或者和其它的脊椎动物一起)有足够的能力将辽东栎坚果捕食或搬运到其它地方分散埋藏或将它们搬运到它们的洞穴中用作漫长冬季的主要食物来源。     

Sequential conflicting selection due to multispecific interactions triggers evolutionary trade-offs in a monocarpic herb

Trade-offs are crucial in understanding phenotypic evolution of organisms. A main source of trade-offs is conflicting selection, a phenomenon very likely in complex multispecific scenarios in which many potential selective agents coexist. The main goal of this study is to investigate the selective trade-offs arising due to conflicting selection on female-fitness components in Erysimum mediohispanicum. I quantified the selection exerted on 10 plant traits by a mutualistic (pollinators) and antagonistic (gall-makers, predispersal and postdispersal seed predators, mammalian herbivores) multispecific assemblage acting sequentially throughout eight selective episodes of the plant, from floral bud to juvenile production. Variation in lifetime female fitness (quantified as number of juveniles) was related mostly to variation in number of flowers, fruit initiation, and seedling establishment. The direction of selection changed among different selective episode for many traits. Most importantly, conflicting selection was frequent in the study system, with half of the phenotypic traits experiencing opposing selection in different selective episodes. Selection at individual life-cycle stages diverged remarkably from selection based on total fitness. Consequently, the evolution of many traits is determined by the relative importance of each episode of selection, with conflicting selection inevitably yielding evolutionary compromises.     

人工埋藏植物种子对鼠类采食下辽东栎坚果存活及萌发率的影响

辽东栎林（Q. liaotungensis)是北京东灵山地区典型地带植被。过去研究表明,鼠类对辽东栎更新有很大影响。本研究主要调查3个问题：（1）采食辽东栎坚果的鼠类种类;（2）将坚果埋藏于土中能否减少鼠类对坚果的采食,并提高出苗率？（3）地表植被是否影响辽东栎的出苗率？1996年9月在北京东灵山地区山地选取5个类型的样地37块,每样地在地表放置10粒坚果,检查地表种子消失率。使用木板夹来调查采食坚果的鼠类种类,诱饵分别为坚果和花生米。于1997年10月,选取5块样地,间隔25 m,每样地在1m²范围内埋50粒坚果,深度5 cm,次年5月调查出苗情况。研究结果表明,大林姬鼠、社鼠和黑线姬鼠为取食坚果的主要种类,取食比例与其丰富度有关。370粒坚果放在地表后,1d、10d、20d的消失率分别达15.1%、78.1%、和 100%。250粒坚果埋入土层后,出苗率达11.2%,苗平均高度为7.8cm,出苗率与地表植被盖度有一定负相关。本研究认为,人工埋藏植物种子可以减少鼠类捕食坚果,并能增加其出苗率。     

Conflicting selection on diaspore traits limits the evolutionary potential of seed dispersal by ants

Conflicts of selection on diaspore traits throughout the dispersal cycle can limit the evolutionary consequences of seed dispersal. However, these conflicts have never been investigated in directed dispersal systems. We explored conflicts of selection through life stages of dispersal in the myrmecochorous herb Helleborus foetidus. Seeds are subject to two contrasting partial selective scenarios. Undispersed seeds are subject to positive directional selection on seed size characters, whereas seeds dispersed are subject to stabilizing selection for size. In both scenarios, seedling establishment determined the magnitude and direction of selection. This does not reflect ant preferences for seed size. However, total selection still depends largely on ant activity, as ants control the relative importance of each selective scenario. We advocate the use of analytical approaches combining multiplicative fitness and microenvironment‐specific selection to more realistically estimate the realized selection on traits functional during several life stages. This approach may be extended to any organism dispersing offspring to different environments.     

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.     

Seed predators exert selection on the subindividual variation of seed size

Subindividual variation among repeated organs in plants constitutes an overlooked level of variation in phenotypic selection studies, despite being a major component of phenotypic variation. Animals that interact with plants could be selective agents on subindividual variation. This study examines selective pressures exerted during post‐dispersal seed predation and germination on the subindividual variation of seed size in hawthorn (Crataegus monogyna). With a seed offering experiment and a germination test, we estimated phenotypic selection differentials for average and subindividual variation of seed size due to seed predation and germination. Seed size affects germination, growth rate and the probability of an individual seed of escaping predation. Longer seeds showed higher germination rates, but this did not result in significant selection on phenotypes of the maternal trees. On the other hand, seed predators avoided wider seeds, and by doing so exerted phenotypic selection on adult average and subindividual variation of seed size. The detected selection on subindividual variation suggests that the levels of phenotypic variation within individual plants may be, at least partly, the adaptive consequence of animal‐mediated selection.     

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.     

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     

Does the spatial variation in selective pressures explain among-site differences in seed mass? A test with Buxus balearica

Positive effects of seed size on germination and survival can be offset by a greater probability of predation or a poorer dispersal of larger seeds. We hypothesized that spatial variation in local selective pressures acting on seed mass may lead to differences in both optimal and observed seed mass among discrete populations. We first examined the variation in seed mass across a total of 14 populations of Buxus balearica; in six of such populations, we further examined the temporal variation by measuring seed mass during three consecutive years. Second, we evaluated seed mass effects on different recruitment phases (seed rain to seedling establishment) in three populations. Lastly, for these three populations, we estimated the resulting overall phenotypic selection on seed mass during recruitment and compared the observed and the predicted optimal seed masses. Most variation (c. 70%) in seed mass occurred among populations, and although we found inter-annual variation in seed mass, the differences in seed mass among populations were consistent over time. Conflicting selective pressures on seed mass appeared during recruitment, and their direction and strength varied among populations, depending on the relative local importance of seed predation vs. germination and establishment. Observed seed mass matched predicted optimal seed mass in two of the three examined populations, suggesting local adaptative responses to the spatial mosaic of selective pressures.     

动物与植物种子更新的关系Ⅱ．动物对种子的捕食、扩散、贮藏及与幼苗建成的关系

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