东灵山地区辽东栎种子库统计

东灵山地区辽东栎成熟林内辽东栎（Ｑｕｅｒｃｕｓ ｌｉａｏｔｕｎｇｅｎｓｉｓ）种群更新记要依赖于萌蘖,而实生苗极少。为探索其原因,我们应用随机样方法进行了辽东栎种子库统计。结果表明：１）种子雨持续时间短、强度大,种子散落总密度为１２３个．ｍ＾－２。２）种子大小在种子雨的不同时期差异很大,大的种子在种子雨高峰中后期所占比例较高。３）土壤种子库存在时间约１００ｄ左右,种子密度最高时为４２．７个．ｍ＾－２     

东灵山地区辽东栎幼苗的建立和空间分布

研究树木实生苗的建立对森林生态系统的保育和恢复具有重要意义,在北京东灵山地区调查了辽东栎(Quercus liaotungensis)实生和萌生幼苗在几种典型的植被类型中的空间分布以及辽东栎一年生实生幼苗在1个林窗梯度上的建立。选择几种典型的植被类型,研究辽东栎实生幼苗和萌生幼苗在森林中的空间分布。结果显示,在不同植被类型中辽东栎实生和萌生幼苗的密度存在空间差异,实生苗密度随幼苗年龄增大逐渐降低,种群的更新主要依靠萌生苗完成。在1个落叶阔叶林中选择1个林窗,在不同梯度(即林窗中间、林窗和树冠连接处,以及树冠下)上,播种辽东栎坚果,第二年秋季调查实生幼苗的2个生长指标和出苗率。结果显示,林窗对一年生实生幼苗的生长高度有影响。幼苗高度在林窗中比在树冠下大;林窗对辽东栎幼苗最长叶片长度和幼苗的出苗率没有影响。说明辽东栎实生幼苗的良好生长需要阳光比较充足的生境条件。本项研究结果建议,对辽东栎种群的更新,应该适当择伐一些较大个体,使森林形成一些林窗或林中空地,以利于辽东栎实生幼苗的建立,这样才能使辽东栎种群通过实生苗进行更好的自然更新。     

间伐强度对黄桥林区辽东栎林结实规律的影响

为了阐明抚育间伐强度对辽东栎林结实规律的影响,以黄土高原南部黄龙山和桥山林区辽东栎中龄林为研究对象,对不同间伐强度(对照、10%、20%、30%)近自然经营抚育间伐5年后的林分密度与结实母树比例、平均单株结实与单位面积林分总结实量、种子在树冠上空间分布特征及种子特征等进行调查分析.结果表明: 辽东栎林结实母树的比例、平均单株结实量及发育种子比例均随着间伐强度的增大而增大,表现为30%>20%>10%>对照;而单位面积林分总结实量在20%强度间伐时达到最大,表现为20%>30%>对照>10%.从辽东栎种子在树冠上的空间分布来看,73.6%的种子集中分布于树冠上层,26.4%分布于下层.树冠阳面结实量占总结实量的65.8%,树冠阴面仅占34.2%,间伐强度对树冠下层、阴面结实量的提升幅度分别大于树冠上层和阳面.辽东栎种子的长径、短径及千粒重均随间伐强度的增大而增大,在30%间伐强度时达到最大.10%强度间伐对辽东栎林结实影响不显著,间伐强度达到20%时,才能显著提高辽东栎种子的数量和品质,继续提高间伐强度到30%时,并不能有效提高其结实,而其林分总结实量却下降.建议在对该林区进行抚育间伐时,应选择20%的间伐强度(保留郁闭度0.7),可有效提高辽东栎种群的结实量和品质.     

啮齿动物和鸟类对东灵山地区辽东栎种子丢失的影响

20 0 0年 8月中旬至 10月上旬 ,在北京东灵山地区小龙门林场选取两块辽东栎分布近似而坡向不同的样地 ,并对样地内种子库与啮齿动物的种群数量变化进行了调查。结果表明 ,两块样地种子雨持续 4 0天左右 ,且种子下落趋势基本一致 ,高峰期都集中在 9月中旬 ,不同坡向的种子产量差异显著。在两样地随机各设置 2 4个种子方形收集器 (0 5m2 )和 2 0个地表样方 (1 0× 0 5m2 )调查种子产量。通过比较收集器内壳斗和种子数量 ,发现二者无显著差异 ,说明鸟类对林冠层种子丢失作用不明显 ;而收集器和地表样方种子数量差异显著 ,表明辽东栎种子库扩散主要由林中啮齿动物完成。标记重捕发现辽东栎林中啮齿动物群落包括大林姬鼠 ,社鼠 ,棕背鼠平和花鼠 ,其中大林姬鼠为优势种 ,占群落的 77.2 %。     

东灵山两个落叶阔叶林中辽东栎种群结构和动态

对北京东灵山地区的两个辽东栎种群的结构和数量动态进行了研究。在研究种群的空间分别格局时,分别将两个种群中的每个辽东栎个体在样地中进行定位。将辽东栎个体按不同尺度大小分为４类,在每一尺度中,按不同格子尺度进行机取样,分别计算Ｍｏｒｉｓｔａ的分布指数,聚块性指数,方差／均值比。用这些指数判断辽东栎不同大小级别以及不同取样尺度上的格局强度。结果表明,两个种群的格局强度都随尺度的不同而变化。５０龄辽东栎林     

秦岭及陕北黄土辽东栎林群落物种移样性特性

应用５种物种多样性测度指标及４种物种－多度分布模型对秦岭和黄土区辽东栎林种子植物物种多样性及木本植物种－多度分布进行了对比研究,结果表明：秦岭辽东栎林无论是群落物种多性还是各层次的物种多样性均高于黄土区辽东栎林。秦岭辽东栎林木栖植物种－多度分布符合对数正态分布而黄土区辽东栎林则符合对数级数分布,分布区气候差异是造成两类辽东栎林物种多样性差异的主要原因。物种－多度分布分析物种多样性分析是一致的。     

秦岭及陕北黄土区辽东栎林群落物种多样性特征

应用５种物种多样性测度指标及４种物种—多度分布模型对秦岭和黄土区辽东栎林种子植物物种多样性及木本植物种—多度分布进行了对比研究,结果表明：秦岭辽东栎林无论是群落物种多样性还是各层次的物种多样性均高于黄土区辽东栎林。秦岭辽东栎林木本植物种—多度分布符合对数正态分布而黄土区辽东栎林则符合对数级数分布,分布区气候差异是造成两类辽东栎林物种多样性差异的主要原因。物种—多度分布分析与物种多样性分析结果是一致的。     

灵空山林区辽东栎(Quercus liaotungensis)种群空间分布格局

为了揭示辽东栎种群空间分布格局的形成机制、种群分布规律和动态变化,采用标准地法,应用偏离指数、Lloyd的平均拥挤度和聚块指数、Morisita指数及均方-区组分析,在5个取样尺度下对不同群落类型中辽东栎(Quercus liaotungensis Koidz.)种群进行了格局分析.结果表明:辽东栎种群的空间分布格局受群落内小环境及辽东栎树种本身的生物学特性的共同影响.气候和立地条件基本一致时,群落内地形起伏不平所形成的小环境以及辽东栎树种强烈的伐桩萌芽更新能力对辽东栎种群的空间分布类型具有显著影响;辽东栎种群的空间分布格局受群落学特征的影响主要体现在群落种类组成和结构上.辽东栎作为共优种,在群落内的分布受制于其它共优种,特别是在油松(Pinus tabulaeformis)的树冠下,辽东栎株数极少;而在无油松分布的辽东栎母树周围,出现辽东栎植株集聚的现象.研究表明辽东栎种群在不同群落类型中多表现为集群分布;随着种群的发育其分布格局由集群分布向随机分布过渡.     

间伐强度对黄桥林区辽东栎生殖构件的影响

为了阐明抚育间伐强度对辽东栎生殖构件的影响,以黄土高原南部黄龙和桥山林区辽东栎中龄林为研究对象,对不同间伐强度(对照、10%、20%、30%)下近自然经营抚育间伐5年后的辽东栎平均单株个体大小、生殖构件产量、比率及空间分布格局特征等进行研究.结果表明: 间伐强度对辽东栎林分平均树高、胸径影响不显著,而与冠幅面积、冠幅体积呈显著正相关;辽东栎标准株的生殖枝、幼果、成熟果实产量随间伐强度的增大而增大,而营养枝、雄花序、雄花、雌花的产量变化不大.当间伐强度达到20%时,生殖枝、幼果、成熟果实产量显著提高,30%间伐下达到最大.生殖枝比例和座果率随间伐强度增大而增大,在20%、30%间伐强度时与对照的差异达到显著水平.约3/4的生殖枝、雌花、雄花集中分布于树冠上层,1/4分布于下层;树冠阳面生殖构件数量占65%,树冠阴面占35%.树冠下层、阴面生殖枝、雌花、雄花序的空间分布比例随着间伐强度的增大而增大;树冠上层、阳面生殖枝、雌花、雄花序分布比例随间伐强度的增大呈递减趋势.因此,选择30%间伐强度(保留郁闭度0.6)最佳,可以大幅提高辽东栎的座果率和成熟率,有效提高结实量和品质,为辽东栎林的采种及天然更新提供保证.     

北京东灵山区两种生境条件下辽东栎幼苗补充与建立的比较

 北京东灵山区的人工油松（Pinus tabulaeformis）林中常常混生有萌生的辽东栎（Quercus wutaishanica）种群。局部地带辽东栎与油松形成混交林,在山脊及山坡上部一些特殊生境辽东栎甚至取代油松林成为优势种。为了探讨北京东灵山区辽东栎林的天然更新机制及辽东栎幼苗在人工油松林中的天然更新,在油松人工林的林下和林缘两种生境条件下将辽东栎种子播种于2 cm深的土中,对辽东栎幼苗的补充和建立进行了对照试验。结果显示,在辽东栎结实丰年,两种生境条件下播种后辽东栎种子丢失差异并不显著,而辽东栎幼苗补充和建立方面的差异极显著。由于森林动物特别是啮齿类动物的活动对辽东栎幼苗建立的影响具有二重性：消耗种子与促进萌发,本试验特别关注辽东栎幼苗子叶丢失现象及其丢失后果。由于辽东栎种子萌根较早,在幼苗出土前其粗大的主根长达10～20 cm,并已初步形成根系,表明已有相当比例的营养物质从子叶转移到根部。因此当幼苗出土后如果仅仅只是子叶的丢失对辽东栎幼苗的成活和生长均无明显的影响。通过研究发现辽东栎幼苗期子叶丢失有3种形式：1）仅丢失子叶,幼苗其它部位未受伤害,多发生在林缘生境中,占发芽坚果的29.69%;2）整个幼苗连根被拖出地面,多发生在林下生境,占62.43%;3）在子叶与根颈的连接处主干被咬断,林缘为50.88%,显著高于林下的8.41%。但只有后两种形式才导致幼苗死亡。导致这些现象的原因是由于林缘土壤腐殖质含量低,比较干燥,土质较坚实,以及幼苗出土前坚果主根生长得较长等缘故,当幼苗遭受动物捕食时不会轻易地被拖出地面,拖走的往往仅仅是子叶。至生长季节结束,林缘样方辽东栎幼苗的成活率接近20%,茎干平均高度7.94 cm,芽数11.24·株-1,叶片平均干重为81.14 mg·株-1,且几乎全部由丢失子叶的幼苗所构成;而林下样方辽东栎幼苗的成活率不足2%,茎干平均高度4.74 cm,芽数7.52·株-1,叶片平均干重仅为42.27 mg·株-1,以未丢失子叶的幼苗为主。试验结果表明,林缘及类似林缘的环境条件更有利于辽东栎的实生更新。     

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.     

浙江天童常绿阔叶林栲树种子雨的时空格局

在浙江天童常绿阔叶林中选择5株栲树,通过种子筐和地表取样,分析了栲树种子雨的时空格局特征.结果表明,栲树种子雨从11月3日到12月16日,持续45d,存在散落的高峰期(11月18日～27日);散落在地表的种子雨主要集中在离母株树干1.5～5.5m区段内,占总数的85.6%,最远不超过8m,空间格局符合二次分布;整个种子雨期间,单株平均落果数为112.8个·m^-2(70个·m^-2～239.5个·m^-2),东南向较多(135.6个·m^-2)、西北向较少(86.8个·m^-2),但无显著差异;种子雨中有活力的坚果占总数的27.3%、未成熟的占25.7%,昆虫幼虫侵害的占42%,松鼠啃食的占5%.     

Utilization of red oak acorns in non-bumper crop year

Utilization of acorns from individual red oaks Quercus rubra Fagaceae) was examined in the tree canopies during fall and from the ground beneath the canopies during winter in a Missouri oak-hickory forest. The goal of the study was to determine whether vertebrates show preference for acorns from individual trees on the basis of acorn or crop characteristics. Seed production, percentage of crop which was viable, maldeveloped, or infested by insects, average seed weight and tannin content of mature, viable acorns were measured for 11 red oak adults. Behavioral observations were conducted in September and October during the peak period of seed fall. Over winter removal was determined by counting the number of acorns removed from 3x3 m plots underneath the trees' canopies between December and mid-April. Canopy removal of acorns was surprisingly low-an estimated average of 52 acorns per tree compared with a mean estimated crop size of approximately 1,200 mature acorns. Over winter, approximately 50% of the acorns were removed from ground plots (range 7% to 93.1%). Number removed corresponded more to the density of nuts rather than to seed and crop characteristics. The low rate of acorn utilization by canopy visitors contrasts with other studies in the literature and may have been due to 1) the fact that this study was done in a relatively homogeneous forest where high density of conspecifics reduces the impact on any one individual or 2) perhaps an unusually low density of birds and mammals.     

Context-dependent post-dispersal predation of acorns in a California oak community

Seed dispersal and predation play important roles in plant life history by contributing to recruitment patterns in the landscape. Mast-seeding – extensive synchronized inter-annual variability in seed production – is known to influence the activity of acorn consumers at source trees, but little is known about its effect on post-dispersal predation. We conducted a planting experiment over three years to investigate the relationship between habitat-level post-dispersal predation and landscape-wide acorn production of three sympatric oak species (Quercus spp.). We measured post-dispersal predation in three oak-dominated habitats – savanna (under Q. lobata), forest edge (under Q. agrifolia), and woodland (under Q. douglasii) – as well as in chaparral and open fields. Overall, landscape-level predation was similarly high among study years, averaging 61.4%. Neither species nor mass of planted acorns affected predation. Habitat had a significant effect on post-dispersal predation risk with acorns disappearing most rapidly in chaparral and least rapidly in woodlands. However, a significant interaction between year and habitat (Z = −4.5, P < 0.001) showed that the hierarchy of predation risk among habitats was inconsistent among years. Using annual acorn census data from local populations of each oak species, we found that predation risk in oak-dominated habitats was significantly and positively related to acorn production of the overstory species (Z = −9.53, P = 0.009). Our findings add to growing evidence that seed dispersal, predation, and regeneration are context-dependent on annual variation in community-level seed production, and we discuss the potential consequences of these dynamics on oak recruitment and animal behavior.     

Acorn production prediction models for five common oak species of the eastern United States

Acorn production varies considerably among oak (Quercus) species, individual trees, years, and locations, which directly affects oak regeneration and populations of wildlife species that depend on acorns for food. Hard mast indices provide a relative ranking and basis for comparison of within- and between-year acorn crop size at a broad scale, but do not provide an estimate of actual acorn yield—the number of acorns that can potentially be produced on a given land area unit based on the species, number, and diameter at breast height (dbh) of oak trees present. We used 10 years of acorn production data from 475 oak trees to develop predictive models of potential average annual hard mast production by five common eastern oak species, based on tree diameter and estimated crown area. We found a weak (R² = 0.08–0.28) relationship between tree dbh and acorn production per unit crown area for most species. The relationship between tree dbh and acorn production per tree was stronger (R² = 0.33–0.57). However, this is because larger-dbh trees generally have larger crowns, not because they have a greater capacity to produce more acorns per unit crown area. Acorn production is highly variable among individual trees. We estimated that dbh of at least 60 dominant or codominant oak trees per species should be randomly sampled to obtain an adequate representation of the range of dbhs (≥12.7 cm dbh) in a given forest area, and achieve precise estimates when using these equations to predict potential acorn production. Our predictive models provide a tool for estimating potential acorn production that land managers and forest planners can apply to oak inventory data to tailor estimates of potential average annual acorn production to different forest management scenarios and multiple spatial scales. © 2011 The Wildlife Society.     

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.     

Multi-trophic effects of ungulate intraguild predation on acorn weevils

Predators and parasitoids may contribute to controlling the population sizes of phytophagous insects, and this has been shown to benefit plants. Phytophagous insects may also be killed by other herbivores (intraguild predation), usually larger-sized vertebrates that ingest insects accidentally while feeding on common food sources. We studied the intraguild predation on acorn weevils by ungulates and assessed the consequences for weevil populations. Infested acorns are prematurely abscised and the weevil larvae finish their development inside the acorns after being dropped. Our results show that weevil larvae were killed by ungulates eating the infested acorns on the ground. Ungulates did not discriminate between infested and sound acorns, and the probability of a larva being incidentally eaten was inversely related to acorn availability. Thus, predation risk was enhanced by the premature drop of infested acorns when acorn availability on the ground was low. Predation rates on infested acorns were much higher where ungulates were present, and acorn infestation rates were significantly lower. However, ungulates did not provide the oaks any net benefit, since the reduction of infestation rates was not enough to compensate for the large amounts of sound acorns eaten by ungulates. Seed predation is usually studied as a progressive loss of seeds by pre- and post-dispersal predators, but the interactions between them are usually not considered. We show that intraguild predation on insects by large ungulates had an effect on the structure of the foraging guild, as the proportion of acorns predated by insects decreased; however, replicating the same experimental design in different ecological scenarios would increase the strength of these results. In conclusion, the present study shows the importance of considering the multi-trophic interactions between seed predators in order to have a complete picture of granivory.     

Seed transfer among bird-dispersed trees and its consequences for post-dispersal seed fate

We investigated seed transfer, i.e. the seed movement away from a source canopy to areas beneath heterospecific canopies, among the ornithochorous tree species Taxus baccata, Ilex aquifolium and Crataegus monogyna in temperate secondary forests in NW Spain, by analysing the composition of multispecific seed rain beneath the canopy of each species, at four sites and for 2 years. To evaluate the consequences on seed fate, we estimated predation by rodents in manipulated seed rains, representing variable levels of relative proportion and total density for combinations of a preferred species paired with a less-preferred species. Seed rain under Taxus canopies was dominated by Taxus seeds, which occurred in low proportion under heterospecific canopies. Ilex seeds dominated the areas under Ilex but accounted for 20–40% of seeds under heterospecific trees. Crataegus seeds were not dominant in any of the microhabitats. The probability of being deposited beneath a heterospecific canopy was much higher for Ilex and Crataegus than for Taxus. The effects of seed rain composition on post-dispersal seed predation were species-specific. Taxus seeds experienced lower predation when occurring in a background of seeds dominated by heterospecific, Ilex or Crataegus, seeds. Crataegus seeds escaped predation more successfully in high-density patches, independently of seed clump composition. Predation on Ilex seeds was independent to both the density and the composition of seed clump. Seed transfer among heterospecific tree species may contribute to shape the template of propagule abundances from which forest will develop, by generating seed combinations favourable to escape from predation.     

Seed predation by mammals and forest dominance by Quercus oleoides,a tropical lowland oak

Summary Quercus oleoides Cham. and Schlecht is an unusual tree in several respects: it is an oak found in neotropical lowland forests, its distribution is not continuous but ratherdivided into many patches of various sizes, and it is a dominant in all the forests in which it occurs, attaining densities far higher than most species of tropical trees. This density pattern is related to the vulnerability of Q. oleoides acorns to predation by mammals. Observations of agoutis, deer, peccaries, squirrels, pocket mice and other seed consumers in Santa Rosa National Park, Costa Rica, showed that these mammals act only as predators, not dispersers, of Q. oleoides acorns. Experiments which involved placing acorns in deciduous forest where Q. oleoides does not occur, demonstrated that, due to high predation rates, the number of acorns produced by an isolated tree is far too low for adults to replace themselves.In oak forest, on the other hand, where the combined acorn crops of many oaks satiate the seed predators, acorn survivorship until germination is high enough to maintain the population. Furthermore, acorn survivorship in oak forest areas is inversely proportional to the apparent mammal density in those areas. Thus the pattern of forest dominance and patchy distribution is related to positively density-dependent acorn survivorship: where Q. oleoides is the forest dominant, it will survive, but if its density falls to the level typical of tropical trees, it will go locally extinct.     

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.