动物搬运与地表覆盖物对辽东栎种子命运的影响

东灵山的辽东栎成熟林内尽管每年有大量种了散落,但实生苗极少,种群天然更新主要依赖于萌蘖。为确定影响种子命运和种子库动态的主要因素,调查统计了不同试验条件下的种子命运。结果发现：（１）动物搬运和取食是影响种子命运和种子库动态的主要因素,在没有该因素的影响下,辽东栎能够完成从种子到幼苗的过程。（３）被啮齿动物贮藏的种子大多被取食,极少存活;（３）地表覆盖物能养活动物搬运、取食的影响,但尚不足导以致新生幼苗出现。     

动物与红松天然更新关系的研究综述

红松球果成熟后,种子自身不能脱落到达适宜的萌发地,因而无法自身完成天然更新而要依赖动物帮助传播种子。捕食红松种子的动物包括啮齿类、鸟类和一些大型兽类。其中,松鼠、星雅和普通币鸟具有分散贮藏的行为,以前两者为重要。这些动物从球果中取出种子,吃掉一部分后,靠颊囊或舌下囊搬运至贮藏地,将种子埋藏在地被物下准备冬、春食物缺乏时再重新捕食。搬运的距离远达数百米至几公里。分散贮藏的贮点数量很大。每个贮点有一至数粒种子。贮点小生境比较适合红松种子的萌发,因此大量的未被重新发现和捕食的贮点的种子在20个月后可能萌发。虽然动物贮藏的生境适合幼苗的萌发,但是不一定适合幼苗的继续生长。由于动物的捕食和贮藏行为,阔叶红松林内红松种子运转途径在4个层次（树上、地面、地被物下和洞穴）进行,最后只有少数种子萌发为幼苗。幼苗的生长过程同样受到动物的捕食和破坏。由于红松结实的丰欠年度变化,动物种群也随之变动。     

动物分散贮食行为对植物种群更新的影响

分散贮食是许多动物取食行为策略的重要组成部分。对以植物种子为主要贮食对象的动物来说,种子内营养物质含量、种子大小以及种子内次生化合物的含量等因素都直接影响动物的贮食行为。动物偏爱贮藏个体较大的种子,大种子多被搬运并分散贮藏在远离种源的地方,而小种子则多被就地取食,以补充动物贮食过程中的能量消耗。贮食动物主要通过空间记忆、特殊路线以及贮藏点周围的直接线索等途径重新获取贮藏点内食物。在重取过程中,一些贮藏点被遗忘,其中的种子成为植物种群更新的潜在种子库。因此,分散贮食动物不仅是种子捕食者,还是种子传播者,它们对植物种子的捕食、搬运和贮藏,影响了植物种子的存活和幼苗的建成,从而在一定程度上影响植物种群的更新、分布。植物种群为了促进种子的传播,在进化过程中逐渐形成了形式多样的适应性策略,降低种子的直接被捕食率,提高种子的被贮藏率。研究动物分散贮食行为对植物种群更新的影响,将有助于理解贮食动物与植物之间的互惠关系,从而认识贮食动物种群在生态系统中的作用,为生物多样性的保护提供科学依据。     

啮齿动物及凋落物存留对天山野果林野生樱桃李种子命运的影响

2010年9月至2013年4月通过设置人工土壤种子库,研究了野生樱桃李土壤种子库的动态及啮齿动物和凋落物覆盖对种子库中种子命运的影响.结果表明: 在有取食动物扰动下,48.3%的种子萌发输出为幼苗,50%的种子被动物搬运或当场取食,4%的种子腐烂.在排除了取食动物干扰的条件下,樱桃李种子形成了短期持久的土壤种子库,3年后依然有部分种子萌发并输出为幼苗.凋落物覆盖和对照处理中,被搬运和当场取食种子的比例均显著低于清除凋落物的裸地.地表凋落物存留能减少动物搬运、取食,但不足以导致新生幼苗的大量出现,而啮齿类动物的搬运或取食是影响野生樱桃李种子命运和种子库动态的主要因素.
     

啮齿动物的贮藏行为与植物种子的扩散

绝大多数啮齿动物一方面取食大量的植物种子和果实,另一方面通过其贮食行为将植物种子和果实搬运到远离母树的地点,即扩散,并将它们分散埋藏在落叶下或浅表的土层中,从而影响种子和果实的时空分布,最后导致幼苗在有利的条件下发生和建成,实现植物更新。啮齿动物与植物种子和果实之间已广泛形成了互惠或协同进化的关系。啮齿动物的贮食行为主要通过以下几个过程对植物种子和果实的扩散产生影响：选择、搬运和埋藏以及随后对种子和幼苗存活和死亡的影响等。本综述了啮齿动物对植物种子和果实贮藏的研究结果,以期为进一步开展啮齿动物的贮食行为在植物种子和果实的扩散中的作用的研究提供参考。     

啮齿动物对秦岭松栎混交林建群种种子扩散格局的影响

森林群落中的啮齿动物对林木种子存在着一定的取食偏好性,这种偏好性会使啮齿动物形成不同的捕食和贮藏策略,从而导致林木种子形成不同的扩散格局。以秦岭中段松栎混交林建群种锐齿槲栎、油松、华山松为研究对象,采用塑料标签标记种子方法,研究啮齿动物对种子传播和扩散格局的影响。结果表明:(1)种子特征是影响啮齿动物对其进行扩散的重要因子,油松种子以其质量小、种皮薄等特点吸引啮齿动物大量捕食,其原地取食率达到83.33%,显著高于华山松和锐齿槲栎种子。(2)锐齿槲栎和华山松种子大部分被搬运一定距离后再被取食和埋藏,其中,扩散距离在1 m以内、1—3 m和5—8 m的锐齿槲栎种子分别占59.12%、18.23%和13.26%,最远扩散距离达12 m处;华山松种子扩散距离主要介于在1—3 m(37.85%)和3—5 m(23.73%),在距离5—8 m(13.56%)、8—10 m(11.86%)和大于10 m(11.30%)区间也有一定的分布,最大扩散距离为12 m。(3)啮齿动物倾向于将锐齿槲栎种子搬运至松林内取食,而将华山松种子搬运至栎林内埋藏,说明不同林分类型及其环境因素对林木种子扩散后的分布格局具有重要影响。(4)3种类型种子被啮齿动物捕食的比例以油松最大(96.90%),锐齿槲栎次之(73.57%),华山松最小(50%);次年调查时,未被捕食的种子大部分已经被取食,仅有极少数锐齿槲栎种子萌发成幼苗(1.67%)。啮齿动物的捕食和贮藏行为对林木种子扩散及其成功更新都至关重要。     

福建闽清福建青冈天然林种子雨和种子库

对福建闽清黄楮林自然保护区的福建青冈(Cyclobalanopsis chungii)天然林的种子雨和土壤种子库进行了观测和分析.结果表明,福建青冈种子雨持续2个月,其高峰期在11月下旬～12月上旬,总量为12.44个m~(-2),在种子散布过程中完好种子、虫蛀种子、败育种子、霉烂种子和萌发种子的比例差异明显,其中虫蛀种子是萌发种子的19.44倍.土壤种子库中虫蛀种子占53.79%,与种子雨相比,完好种子和萌发种子数量分别减少2.15个m~(-2)和0.20个m~(-2).土壤种子库种子存活率仅13.51%,动物捕食率达45.90%,说明动物的捕食和搬运是福建青冈种子缺失的原因之一.福建青冈天然林土壤种子库有69种植物种子,但种子数量较少,且分布不均匀,种子库中78.42%种子分布在2～5 cm土层中.福建青冈天然林中实生幼苗少与福建青冈种子本身特性及其生长环境密切相关,福建青冈种子发育成熟后因动物侵扰、虫蛀、霉烂和败育等情况发生,难以在土壤中长期保存,以致福建青冈林分天然更新严重受阻.     

不同贮藏温度下大别山五针松花粉活力的变化

大别山五针松( Pinus dabeshanensis C. Y. Cheng et Y. W. Law)为松科( Pinaceae)松属( Pinus Linn.)植物,自然种群数量极少,目前仅发现在安徽省岳西县大王沟海拔900~1300 m阴坡和半阴坡有相对集中的分布,种群规模计200余株,且多为成年个体,林下幼苗极少,自然更新困难,为中国特有珍稀树种之一[1]。大别山五针松常在每年3月份至4月份形成花芽,5月中下旬花粉成熟并散发,花粉具气囊,此时雌球花张开接受传粉,球果翌年9月成熟[2]。据作者近年的调查,大别山五针松种子败育率较高,且种子质量较低,这也是该种类濒危的重要原因之一。     

啮齿动物对六盘山区辽东栎、野李和华山松种子的取食和搬运

为了阐明不同种子投放方式和覆盖处理对啮齿动物取食和搬运/贮藏种子行为的影响,2010年春季,我们在宁夏六盘山区的辽东栎幼林,用塑料标签标记法研究了不同种子投放方式(单独投放、两两组合投放和混合投放)和覆盖处理(对照、凋落物覆盖和土壤覆盖)下啮齿动物对辽东栎、野李和华山松种子的取食和搬运特点。结果表明:(1)不同物种、不同种子投放方式和覆盖处理对种子的就地取食率、搬运后的取食率和贮藏率均具有显著影响,种子较大或具有厚而坚硬的种皮(内果皮)、组合和混合投放及覆盖处理均不同程度地促进啮齿动物对种子的搬运;(2)3种植物的种子被搬运后取食和贮藏的平均距离范围均在5.5m以内,而且种子被搬运后贮藏的距离均大于搬运后取食的距离,其中华山松种子被搬运后取食的距离显著大于辽东栎和野李种子,但搬运后的贮藏距离显著小于后两者。(3)种子被搬运后单个种子的取食点和贮藏点占多数,其他大小的取食点和贮藏点较少,种子被搬运后以土壤埋藏为主要的贮藏方式。结果表明,种子的种皮特征以及种子在群落中的分布方式和存在状态,可能通过延长啮齿动物处理的时间而增大啮齿动物的被捕食风险,使动物改变对种子的取食和搬运行为,进而决定了种子的命运。     

濒危植物银杉的结实特性及动物对果实的危害性

银杉结实量的年际波动很大,具有明显的大小年之分,一般在２～３个小年之后为１～２个大年。银杉母株的结实量在不同的母树间差别很大,平均单株的球果数仅为６６个。球果结实量在树冠南北侧有差异,南侧球果数占球果总数的６０％;球果结实量的垂直分异明显,树冠中、上部的球果数占总量的９０％以上。松鼠对树上尚未完全成熟的银杉果实的取食危害是长期性的,对银杉种子种群造成严重破坏。以１９９５年为例,被啃食的球果数占球果总数的比重可达１５％左右。试验表明,土壤种子库中地表的银杉种子几乎全部被动物取食,种子埋藏深度增加有利于逃避动物的取食。     

Silvicultural characteristics and natural regeneration of Pinus brutia Ten. — a review

Pinus brutia Ten. is a characteristic species of the eastern Mediterranean. P. brutia and P. halepensis Mill. are distinct species; but P. eldarica, P. stankewiczii and P. pityusa are subspecies of P. brutia. Fire is the major disturbance in P. brutia forests, and several adaptations generally contribute to post-fire regeneration of P. brutia; however, P. brutia can also naturally regenerate without fire. Flowering occurs between March and May. Seed distribution occurs throughout the year, with maximum dispersal in August and the great majority of seeds dispersing within the period July-December.P. brutia retains some closed cones and so establishes a natural canopy seed bank. Seed germination occurs in the rainy season with two frequency peaks: the major one in spring and a minor one in autumn. P. brutia seedlings develop rapidly-growing tap roots. Shelterwood, clearcutting and strip clearcut methods can successfully be applied to P. brutia forests, depending on local conditions. Density of P. brutia seedlings was greater and seedlings were more vigorous when the natural regeneration methods were combined with prescribed fire. Clearcutting methods combined with laying cone bearing branches on the surface of the soil and additional seeding increases regeneration success. Ground litter of 2–4 cm creates a medium that reduces evapotranspiration, delays growth of competing vegetation, and increases seedling survival. Successful regeneration must be established by the end of first vegetation period.     

文峪河上游华北落叶松林的种子雨、种子库与幼苗更新

华北落叶松林下更新不良,为探究其制约因素,开展了山西省文峪河上游5个华北落叶松林分的种子雨、土壤种子库和幼苗更新的研究。结果表明:(1)华北落叶松种子主要集中于9—10月散落。2011年为华北落叶松种子丰年:种子产量高,种子雨密度达(961.93±377.40)粒/m2;种子质量高,完整种子占(89.31±16.13)%。2012年为种子平年,种子产量低,种子雨密度为(252.73±115.12)粒/m2。华北落叶松种子雨主要源于毗邻树木,华北落叶松纯林和落叶松云杉林的种子雨密度显著高于其他3个针阔混交林。(2)土壤种子库主要由上年种子雨组成,2012年4月的土壤种子库密度为(695.18±297.23)粒/m2,完整种子占(59.73±9.56)%。种子自然萌发前,约(78.98±24.76)粒/m2具发芽力,基本可满足更新需要。但种子活力保持期少于2 a,只能形成短期持久土壤种子库。(3)华北落叶松更新不良,种子年后仍难以实现幼苗建成,当年生幼苗的出现频度平均为1.6%,且林下难以存活。幼苗发生与种子储量关联性不强,种源条件不是制约华北落叶松更新的主要因素。     

格氏栲天然林林窗和林下种子散布及幼苗更新研究

以格氏栲(Castanopsis kawakamii)天然林为研究对象,探讨了林窗和林下格氏栲种子雨、种子库的分布特征及幼苗更新状况。结果表明:林窗种子雨总量和完好种子密度高于林下,未成熟种子比例低于林下;林窗和林下种子雨高峰期掉落数量分别占种子雨总量的77.13%和74.5%;林窗种子库储量低于林下,种子库中以全食或捡拾种子比例最高,其中种子库储量垂直分布表现为枯落物层(约占2/3)>腐殖质层(0～5 cm)(约占1/3)>心土层(5～10 cm)(小于1%),以格氏栲种子占绝对优势;格氏栲从种子到幼苗的转化率低,林窗中格氏栲完好种子密度与幼苗密度均高于林下。林窗微生境提高了种子散布过程中格氏栲成熟种子的密度和比例,有利于促进格氏栲的幼苗更新,表明林窗在格氏栲种群恢复过程中扮演着重要角色。     

海南霸王岭陆均松种子雨和土壤种子库特征

为了解陆均松(Dacrydium pierrei)的自然更新能力,对海南霸王岭陆均松进行为期3年的种子雨、种子库及幼苗幼树观测,并分析其与环境因子的关系。结果表明,陆均松种子雨和种子库有效种子数量太少并存在季节性差异;其结实存在大小年现象;种子雨的扩散密度随着距母树距离的增加而逐渐减少;土壤种子库种子数量在枯枝落叶层所占比例最大,但有活力种子数量极少。幼苗幼树适宜生存的海拔范围为800~1 400 m,经度、纬度、坡度与幼苗幼树分布有明显相关性。因此,选择适宜季节采摘种子,提高种源数量和质量,清除林下枯枝落叶,增加种子萌发能力等以减少陆均松更新的限制条件的人工抚育措施应适当考虑。     

The role of seed dispersal and seedling traits in colonization and coexistence ofSalix species in a seasonally flooded habitat

Regeneration traits of six co-occurringSalix species were studied on a floodplain of the Sorachi River, central Hokkaido, Japan, and their colonization success and coexistence in a local habitat were discussed. MixedSalix communities contained sixSalix species; dominant:S. sachalinensis; four subordinates:S. rorida, S. pet-susu, S. miyabeana andS. subfragilis; rare species:S. jessoensis. Their phenology, falling velocity and longevity of seeds, and the effects of microtopography and soil texture on seedling establishment were studied. The sixSalix species had overlapped seed dispersal periods that coincided with the decrease of water level after a predictable spring flood. This coincidence was crucial for the colonization success because the seedlings were established on wet soils left by the decreasing water level. They showed two types of regeneration trait, specialization and generalization.S. rorida andS. subfragilis showed contrasting regeneration traits; early vs. late seed dispersal, large vs. small seeds, seedling distribution on coarse vs. fine soils, respectively. These two species rarely co-occurred. On the other hand, the dominantS. sachalinensis had an intermediate seed size and dispersal timing, and a wide range of seedling distribution from coarse to fine soils. These results revealed that flooding seasonality influenced the colonization success together with the regeneration traits ofSalix species, and that coexistence of theSalix species was facilitated primarily by regeneration niche separation related to flooding seasonality and soil heterogeneity.     

Cones structure and seed traits of four species of large‐seeded pines: Adaptation to animal‐mediated dispersal

Seed dispersal selection pressures may cause morphological differences in cone structure and seed traits of large‐seeded pine trees. We investigated the cone, seed, and scale traits of four species of animal‐dispersed pine trees to explore the adaptations of morphological structures to different dispersers. The four focal pines analyzed in this study were Chinese white pine (Pinus armandi), Korean pine (P. koraiensis), Siberian dwarf pine (P. pumila), and Dabieshan white pine (P. dabeshanensis). There are significant differences in the traits of the cones and seeds of these four animal‐dispersed pines. The scales of Korean pine and Siberian dwarf pine are somewhat opened after cone maturity, the seeds are closely combined with scales, and the seed coat and scales are thick. The cones of Chinese white pine and Dabieshan white pine are open after ripening, the seeds fall easily from the cones, and the seed coat and seed scales are relatively thin. The results showed that the cone structure of Chinese white pine is similar to that of Dabieshan white pine, whereas Korean pine and Siberian dwarf pine are significantly different from the other two pines and vary significantly from each other. This suggests that species with similar seed dispersal strategies exhibit similar morphological adaptions. Accordingly, we predicted three possible seed dispersal paradigms for animal‐dispersed pines: the first, as represented by Chinese white pine and Dabieshan white pine, relies upon small forest rodents for seed dispersal; the second, represented by Korean pine, relies primarily on birds and squirrels to disperse the seeds; and the third, represented by Siberian dwarf pine, relies primarily on birds for seed dispersal. Our study highlights the significance of animal seed dispersal in shaping cone morphology, and our predictions provide a theoretical framework for research investigating the coevolution of large‐seeded pines and their seed dispersers.     

The importance of seed trees in the dioecious conifer Pilgerodendron uviferum for passive restoration of fire disturbed southern bog forests

Biological legacies are important for ecosystem recovery following disturbance as demonstrated by studies in the northern hemisphere. Southern bog forests dominated by the conifer Pilgerodendron uviferum in Northern Patagonia are a typical case of an ecosystem with low resilience to disturbance by fire, which kills most trees and seeds, on which the species depends for regeneration. In this study, we hypothesize that the natural recovery of P. uviferum populations in burned areas is limited by seed availability and this limitation may be exacerbated by the dioecy of the species. Using a multi‐scaled approach, we quantified the seed dissemination potential from P. uviferum seed trees, assessed the suitability of substrates for the germination of seedlings, and finally analysed the spatial distribution of seed trees of the species at the landscape level. Our results indicate that 70 years after a fire on Chiloé Island (43°S), natural regeneration from seed trees can assist the recovery of P. uviferum populations following large‐scale fire disturbance, but their effect is limited at a landscape level owing to a low number of reproductive female trees (0.3 trees ha^?1) and limited seed dispersal (<20 m). In this context, a mixed passive‐active restoration approach that takes into account the spatial pattern and sex of seed trees could be the most effective and efficient option to restore not only P. uviferum forests in North Patagonia, but also other heavily disturbed forests with few remnant seed trees, in particular of dioecious species.     

The role of birds and mammals in Korean pine (Pinus koraiensis) regeneration dynamics

This study examined the interrelationships of the fall seed-foraging guild with Korean pine (Pinus koraiensis) and its regeneration. The study took place in old-growth and secondary-growth forests in Northeastern China. Foraging behavior, seed transportation distance, and cache location of various birds and mammals were observed. Regeneration characteristics of Korean pine were also analyzed using plots in various vegetation cover types, successional stages, and topographical situations. Ten species of vertebrates were able to harvest seed from the closed-cone Korean pine. Of these, the Eurasian nutcracker, Eurasian nuthatch, red squirrel, and Siberian chipmunk were found to be potential seed dispersal agents. The nutcracker was the most important dispersal agent, easily acquiring seed with its large pointed bill, carrying up to 62 seeds in one trip, carrying seed at least 4 km, and placing seed in a variety of sites 2.5–3 cm deep in the soil. The Eurasian nuthatch carried a single seed per trip at distances less than 50 m, needed nutcrackers to open the cones and expose seed before they could acquire seed, and occasionally cached seed in the soil. Red squirrels were uncommon visitors to the tree tops of Korean pine, carried cones shorter distances than nutcrackers, and were only found under forest canopies. Human harvest of cones by knocking off branches also affected squirrel behavior and reduced future cone crops. Siberian chipmunks also collected seed from cones in trees and appeared to transport seed less than 50 m. Six other species were observed in this study successfully harvesting seed from cones but were not potential seed dispersers. Natural seedling establishment was found to be over 1000 seedlings/ha except on old-growth pine-hardwood sites. Squirrels were commonest here, but few seedlings survived past the 2nd year due to the intense shading. Second-growth forest types, including aPicea plantation where nutcrackers cached seed daily, and an old-growth pine-hardwood selective-harvest site, had the greatest regeneration. In conclusion, most natural regeneration of Korean pine in this part of its range is due primarily to the Eurasian nutcracker. Nutcrackers can aid forest managers in reaching desired stocking levels after disturbance, as well as a more natural-appearing forest. Squirrels, chipmunks, and nuthatches are minor seedling establishment agents. Korean pine seed is an important food source used by at least 22 species of forest wildlife.     

祁连山排露沟流域青海云杉天然更新与土壤水热的关系

青海云杉是祁连山区森林生态系统的建群种,其天然更新对维持祁连山森林生态系统功能具有重要意义。以祁连山排露沟流域青海云杉林天然更新为研究对象,于2021年6-9月测定海拔2700-3300m的0-40cm土层土壤温度和水分数据,并于7月中旬对样地天然更新进行每木检尺,将天然更新划分为幼苗(第一龄级)阶段和幼树(第二龄级)阶段,通过方差分析探究不同海拔更新苗胸径、株高的差异,再进一步建立回归模型拟合更新苗胸径、株高与各层土壤水热的关系,探讨更新苗胸径、株高对土壤水热的响应。结果表明:(1)更新苗胸径、株高随海拔升高而降低,海拔2700m幼苗胸径与3300m幼苗胸径存在显著性差异(P<0.05),海拔2700m、2800m幼苗、幼树株高与3300m幼苗、幼树株高存在显著性差异(P<0.05),幼树胸径在各海拔均不存在显著性差异,表明更新苗胸径对海拔的敏感性要低于更新苗株高。(2)更新苗胸径、株高与土壤温度呈正相关,与土壤水分呈负相关,幼苗胸径与20cm土壤温度和土壤水分的相关系数最大,幼树胸径与40cm土壤温度和土壤水分相关系数最大,幼树株高与40cm土壤温度及20cm土壤水分相关系数最大。(3)回归模型显示幼苗胸径和株高对土壤水热的利用模式相同,而幼树胸径和株高对土壤水热的利用模式存在差异。(4)青海云杉天然更新随着龄级的增加,影响生长的主要因素由土壤温度逐渐变为土壤水分,更新苗胸径对土壤水热的响应也随着龄级的增加在逐渐减弱。     

Site quality effects on post‐fire regeneration of Pinus brutia forest on a Greek island

Abstract. Natural stands of Pinus brutia were compared to burned areas after a great fire in 1985 on the island of Thasos, North Greece, in relation to site quality effects on the success of natural regeneration and understorey vegetation. The number of seedlings in the natural stands of P. brutia was 1124/ha against 3188/ha in the burned area. In both cases natural regeneration was successful. Under the canopy of P. brutia forests the predominant shrubs were Quercus coccifera and vPhillyrea latifolia. A high percentage (up to 40%) of the total area was covered by these species. In the burned areas, 12 yr after the wildfire of 1985, Cistus incanus and C. salvifolius played a significant role in the survival of P. brutia seedlings. These species had covered the burned area long before the pine seed germination.