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。     

凉水自然保护区松鼠贮藏红松种子距离的初步测量与分析

2006年10月,应用线牌标记红松(Pinus koreansis)球果,采用人工放置实验和定点观察相结合的方法,对松鼠(Sciurus vulgarls)贮藏红松种子距离进行了初步研究与分析.结果表明,不同放置点的红松球果堆被松鼠发现并捕食的时间不一致,放置在人为干扰较强地点(旅游步道边)的红松球果最后被搬运走,但各放置点的红松球果均在放置10 h内全部消失.新球果(2006年)和旧球果(2005年)均被松鼠捕食并贮藏,但松鼠优先选择搬运新球果.实验共放置人工标记的球果110枚,当场被取食的球果8枚,其中花鼠(Eutamias sibiricus)当场取食4枚新球果,松鼠当场取食4枚,均为旧球果;其余102枚球果被搬运并贮藏.以放置点为圆心,以300 m为搜索半径,共回收标志球果47枚,回收率为46.1%.     

啮齿动物的分散贮食行为

食物贮藏是许多动物重要的适应性行为,分散贮藏的食物以植物种子为主。每个贮藏点贮藏数量不等的食物项目。啮齿动物分散贮藏食物之后,可降低食物被其他个体获取的机率,提高对食物资源的控制能力,最终有利于自身的生存和繁殖成功。植物种子被贮藏之后,可减少非贮食鼠类对种子的取食。同时,合适的微生境和埋藏有利于种子萌发、幼苗建成和植物的更新;使植物的分布区得以扩展。探讨啮齿动物的分散贮食行为,能够更好地理解食物贮藏在啮齿动物生活史中的作用,进一步认识鼠类和植物的相互关系以及不同啮齿动物在群落形成中的潜在作用。本综述了啮齿动物分散贮食的研究进展,并提出今后工作中的几点建议。     

动物在大别山五针松种群天然更新中的作用

大别山五针松(Pinus dabeshanensis)为我国特有树种,其天然植株较少且分布范围狭窄,种群更新困难,已被列为国家二级重点保护野生植物。分别于2015年和2016年的10—12月,在目前已知最大天然种群所在地安徽岳西县河图镇大王沟,研究了大别山五针松球果与种子特征、种子雨与土壤种子库、动物对种子的取食和搬运,幼苗分布格局及其与鼠穴分布的关系,以期探明动物在其天然更新中的作用,分析其天然更新不力的原因。结果显示:大别山五针松种子败育率较高,阳坡个体球果发育情况好于阴坡;种子成熟期间没有明显的种子雨,土壤种子库也未调查到完整种子。共记录到母树与球果的访问动物6目11科16种,其中7种动物确定取食种子;地面种子摆放实验显示超过95%以上的种子被啮齿动物捕食或搬运至他处取食或贮藏,不同的摆放处理对啮齿动物的捕食、搬运没有显著影响;小林姬鼠(Apodemus sylvaticus)盗食现象严重,埋藏实验中人工贮点当夜被发现的概率在90%左右,岩松鼠(Sciurotamias davidianus)和小林姬鼠是其种子主要捕食者。研究地的大别山五针松种群于2015年和2016年分别新增一年生幼苗5株和7株,这表明大别山五针松在当地存在天然更新。幼苗多单独生长在母树周围,点格局分析显示在0—0.6 m的尺度范围内呈随机分布;在0.6 m尺度呈聚集分布。大别山五针松幼苗在小林姬鼠巢穴周边分布,其更新格局受到小林姬鼠贮点位置分布情况影响,小林姬鼠极有很可能为大别山五针松的传播者,啮齿动物对大别山五针松种子的捕食与搬运影响了大别山五针松的天然种群更新。     

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%)。     

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。     

红松结实量与松果可利用量对松鼠和花鼠贮食行为的影响

植物通过每隔几年产生大量种子的丰年策略提高贮食动物传播种子效率,但人为采摘活动降低了种子的可利用量,从而影响贮食动物行为及种群动态。为研究红松球果采摘如何通过降低松果可利用量而影响贮食动物行为,我们基于黑龙江凉水自然保护区2003-2012年红松结实量和采摘量的变化,比较分析了结实大小年间松鼠贮点大小、贮点深度、贮食密度和贮藏量、松鼠花鼠种群以及2010年和2011年花鼠洞穴贮藏量的差异。结果显示：结实大年松鼠的平均贮点大小显著高于小年,大贮点比例增加,贮食密度和贮藏量也明显高于结实小年,但随着结实量的增大,球果采摘量增加,使松果可利用量减少,由此结实大年2011年松鼠贮食密度并未随结实量增加而增加,反而低于2003年和2008年,松鼠遇见率也没有在该结实大年有所增长。而花鼠种群和洞穴松籽贮藏量在结实大年和小年间没有显著差异。研究表明红松球果采摘对松果可利用量和松鼠贮食行为有较大影响,应合理确定大年的采摘量以保证贮食动物的食物丰度,维系红松生态系统健康。     

星鸦的贮食行为及其对红松种子的传播作用

1994－1998年秋在小兴安岭凉水自然保护区对星鸦（Nucifraga caryocatact）的贮食行为及具对红松（Pinus koraiensis）种子的传播作用进行了研究,星鸦在秋季至翌年早春主要以红松种子为食,偶食浆果。秋季种子成熟时,星鸦在取食的同时储藏大量种子作为冬季和早春的食物,这些种子中未被完全吃掉的遗留部分在第三年春天即可发芽,在地上取食部分种子后,星鸦通常储存30－50粒于舌下囊中,飞行2－5km后,将种子埋藏在土层下2.5-3.5cm深处,每个贮点从为2－4粒,贮点生境主要有8种：林缘（路边）、保护区周围空阔地、人工落叶松林、人工自冷杉林、人工红松林、人工红皮云杉林、天然更新的杨桦林和天然原始红松阔叶林（母树林）。星鸦每天至少搬运种子10次,达400粒。一个贮藏季节一只星鸦可贮藏至少16000粒种子,对红松1年生和2年生幼苗调查表明,幼苗在上述8种生境中均有分布,并且在人工落叶林、天然更新杨桦林和母树林下有幼树分布,因此星雅能有效地扩散红松种子,有利于红松的天然更新。     

围栏条件下捕食风险对岩松鼠贮藏核桃种子行为的影响

面对捕食风险,贮藏植物种子的啮齿动物如何通过调整其贮藏行为,权衡捕食风险和贮食效益,需要深入研究。2006 年9 月,在北京东灵山地区,以黄鼬作为捕食风险源,在半自然围栏内比较了有无捕食风险源时,岩松鼠贮藏的核桃种子的状态,以及埋藏种子在实验围栏内的分布、微生境和搬运距离,以探讨捕食风险对岩松鼠贮藏种子行为的影响。结果表明,捕食风险源存在时,搬运和埋藏种子的比例都显著减少,弃置地表种子的比例无显著差异;埋藏于高风险区的种子比例减少,中风险区和低风险区的埋藏种子比例增加,但差异都不显著;位于裸地和草下的埋藏种子比例减少,位于墙基的埋藏种子比例增加,但差异都不显著;此外,埋藏种子的搬运距离也没有显著差异。可见,捕食风险源存在时,岩松鼠显著减少了搬运和埋藏种子的比例,但对埋藏种子时的埋藏区域和微生境选择,以及搬运距离无显著影响。差异不显著可能与围栏效应有关。     

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.
     

凉水自然保护区松鼠（Sciurus vulgaris）贮食生境选择

2003年10月~2004年4月,在黑龙江省小兴安岭凉水国家级自然保护区,应用跟踪观察法、样线调查法、样方调查法等研究方法,采用Vanderploeg和scavia选择系数Wi和选择指数Ei作为衡量指标,对松鼠贮食红松种子生境选择进行了研究。研究表明,松鼠贮食对生境具有选择性:以主要林型为划分标准,松鼠偏爱贮食生境依次为:原始红松林,云杉林,人工云杉林,针叶混交林,人工落叶松林,针阔混交林,阔叶混交林,次生白桦林和人工红松林。在原始红松林内,松鼠对微生境的利用存在选择性差异:松鼠对郁闭良好(郁闭度大于0.6)、灌丛密度中等、倒木和枯立木密度中等(1~5个)、盗食动物活动较弱(大林姬鼠Apodemus peninsulae为主要盗食动物)的原始红松林生境表现出较强的选择性(Ei>0.2);而对坡度小于5°、基质为霉菌土、倒木和枯立木密度较高(大于5个)、草本盖度较高、盗食动物活动中等(花鼠Eutamias sibiricus为主要盗食动物)的原始红松林生境表现出较强的负选择性(Ei<-0.2),对阴坡表现出强烈的回避性(Ei=-0.5368)。     

Hoarding habitat selection of squirrels (Sciurus vulgaris) in Liangshui Nature Reserve, China

This study was conducted during October 2003–April 2004 in the Liangshui National Nature Reserve of the Xiao Xing'an Mountains of Northeast China. Results showed that hoarding behavior of squirrels exhibited selectivity. The preference order of hoarding habitat selection of squirrels is as follows: original Korean pine forest, secondary natural fir forest, artificial fir forest, mix-conifer leaf forest, artificial fallen leaves pine forest, mix-conifer-broadleaf forest, mix-broadleaf forest birch forest, and artificial Korean pine forest. Compared with the existing results, using the cache spots as an index, the order of habitat selection changed, revealing that this research should include more factors, such as pilferage from other animals and secondary dispersal by the squirrels or other species. The Vanderploeg and Scavia selectivity indexes, Wi and Ei, were used to evaluate the use of microhabitat by squirrels in the original Korean pine forest. Results indicated that the squirrels exhibited a significant microhabitat utilization pattern: (1) Squirrels prefer to use the microhabitat in original Korean pine forest with high canopies, the medium shrubby density, and the medium stub density, where pilferage animals are few (Ei > 0.2); (2) Squirrels did not prefer to use the microhabitat in the original Korean pine forest with a low slope degree, high density of fallen logs and stumps, a high herbage coverage, and an abundance of pilferage animals (Ei < ?0.2), and they showed an aversion for a shaded slope (Ei = ?0.5368). The selectivity of microhabitat utilization for the hoarding behavior of squirrels plays an important role in determining the spatial pattern of Korean pine seedlings.     

松果采摘对长白山自然保护区红松天然更新的影响

在野外调查研究基础上,结合20世纪70年代未受干扰的红松天然林地面红松种子贮藏情况,分析了长白山自然保护区松果采摘后,红松林地面红松种源贮藏和分布变化,根据红松更新过程与动物的关系。探讨了松果采摘对红松更新条件的影响．结果表明,松果采摘后。地面红松种子主要分布在地被物下层且大多呈单粒状分布,其贮藏量1582～2640粒·hm^-2。仅为20世纪70年代的0．3％～0．5％。废种子比率67．8％～86．2％。单粒状占总簇数的46．8％～77．1％,对红松天然更新起重要作用的动物已减少或消失。表明松果采摘是长白山自然保护区红松天然更新的最大障碍之一．     

Seed dispersal of the Korean pine,Pinus koraiensis,by the red squirrel,Sciurus vulgaris

Seed hoarding behavior of the red squirrel,Sciurus vulgaris, was studied in relation to the amount of dispersed seeds of the Korean pine,Pinus koraiensis, and the distribution of its seedlings. After removing a cone from a tree, squirrels sat on the ground and ripped off its cone scales before transporting it. A mean of 3.2 seeds were scatter-hoarded per hole. Of 7.7×10⁴ mature seeds produced in a 0.21 ha planted Korean pine forest, 22% were estimated to be directly eaten by four squirrels, 9% were hoarded by them in the pine forest and 65% were cached outside the forest. Squirrels rediscovered hoarded seeds frequently, until the ground was covered with snow, during the period from snow fall until seed germination the next spring, few hoarded seeds were utilized. Korean pine seedlings were found up to 600 m from their mother trees. Scatter-hoarding by squirrels extensively contributes to seed dispersal to places suitable for the regeneration of the Korean pine. The large size of the cone, the absciss-layer at the cone penduncle, the infrequent dehiscence of cone scales, the large and wingless seeds, and the thick seed-coats have probably all been specialized to facilitate utilization by GenusSciurus.     

凉水自然保护区松鼠和星鸦贮食生境选择差异

2003 年9 月30 日～2005 年10 月8 日,在黑龙江凉水国家级自然保护区,应用样方调查法, 采用Vanderploeg和Scavia 选择系数Wi和选择指数Ei作为衡量指标,对松鼠和星鸦贮食生境选择进行了研究。结果表明, 二者贮食生境选择优先顺序略有不同,松鼠偏爱的贮食生境依次是: 云杉林、原始红松林、人工红松林、针阔混交林、人工云杉林、白桦林、针叶混交林、人工落叶松林、阔叶混交林、冷杉林和其它。星鸦对贮食生境的偏爱程度依次为: 人工红松林、原始红松林、云杉林、人工云杉林、针阔混交林、阔叶混交林、白桦林、针叶混交林、人工落叶松林、冷杉林和其它。在对贮食微生境因子的选择利用上, 二者大致相同,只是在对优势灌丛的选择上略有差异, 松鼠优先选择在狗枣猕猴桃优势灌丛内贮藏红松种子, 而星鸦优先选择在刺五加优势灌丛内贮食。松鼠和星鸦贮食生境选择的差异将对随后的红松天然更新过程产生不同的影响。     

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.     

长白山阔叶红松林带内杨桦林动态模拟

利用改进的ZELIG .CBA模型模拟了长白山杨桦林的动态变化过程 .根据红松种源的存在与否 ,将模拟分为两种方案 ,结果表明 ,红松参加演替与否决定了山杨、白桦退出林分时间的早晚 ;红松种源存在时 ,演替的最终结果是阔叶红松林 ;无红松种源时 ,演替的最终结果是阔叶混交林 ,两种林分的生物量和蓄积量相差很大     

Weak olfaction increases seed scatter‐hoarding by Siberian chipmunks: implication in shaping plant–animal interactions

Scatter‐hoarding of seeds by animals plays an essential role in seed dispersal of plants and in shaping plant–animal interactions in forest ecosystems, but the function of scatter‐hoarding behavior is still unclear. We hypothesize that weak olfactory cues between seeds and scatter‐hoarding animals would increase scatter‐hoarding. Using a rodent–plant system of Siberian chipmunks Tamias sibiricus and Korean pines Pinus koraiensis, we tested the effects on seed scatter‐hoarding intensity by measuring and modifying the seed odor intensities and the abilities of the animals to detect seed odor. Siberian chipmunks seemed to scatter‐hoard more seeds with weaker odor signals, and Siberian chipmunks with reduced olfactory ability scatter‐hoarded more seeds, supporting our hypothesis. Our studies suggest that olfaction may have played an important role in the evolution of the strength of seed odor and scatter‐hoarding behavior of animals, and in shaping plant–animal interactions.     

The effects of hoarding habitat selection of Eurasian red squirrels (Sciurus vulgaris) on natural regeneration of the Korean pines

Cone-cores discarded by Eurasian red squirrels were used to study the habitat selection of Korean pine-seeds hoarding, in forest patch Nos. 16 and 19 in Liangshui Nature Reserve, China. Ten transects with a total length of 15 km were uniformly set, and data from 343 valid samples were collected in a 369 hm² area. One hundred and eighty four were hoarding samples which were determined according to the cluster analysis based on the number of the cone-cores, while the other 159 were control samples. The principal component analysis, using 11 habitat factors, suggested that the distance from Korean pine forest, forest type, number of Korean pine seedlings, density and type of bush significantly influenced the habitat selection of hoarding by Eurasian red squirrels. The results of Bailey’s method indicated that the squirrels showed (1) preference for natural coniferous forest, natural fir and spruce forest and planted spruce forest; (2) avoidance of planted Korean pine forest and planted larch forest; and (3) random use of natural Korean pine forest. Moreover the distance from the Korean pines in the range of 150–600 m showed no effect on the habitat selection of hoarding by the Eurasian red squirrels. More than 50% of the cone-cores were discarded in either fringe or gap of the Korean pine forest with more cone-cores found at <300 m than at 300 m away (One-Way ANOVA; df = 3, 183, F = 5.76, p = 0.0009). This demonstrated that the Eurasian red squirrels could take the cone-cores out of the Korean pine forest. The density of bushes in samples of hoarding area was significantly lower than that in control samples (Kruskal–Wallis test; df = 1, χ² = 83.99, p < 0.0001). The number of the Korean pine seedlings in samples of hoarding area was significantly higher than that in the control samples (Kruskal–Wallis test; df = 1, χ² = 104.13, p < 0.0001). This illustrated that the hoarding habitat favoured the germination of the Korean pine seedlings. In conclusion the behavior of hoarding Korean pine seeds by the Eurasian red squirrels can promote the regeneration and dispersal of the Korean pines.