岩松鼠贮藏种子的空间记忆研究

通过在岩松鼠自身埋藏种子附近选择相同生境人工埋藏种子的方法,研究岩松鼠对自身埋藏种子和人工埋藏种子的找回率有无差异,从而部分揭示记忆如何在岩松鼠找回贮藏种子过程中发挥作用。数据的统计分析采用SPSS for Windows(version 19.0)进行。结果显示:1)岩松鼠喜欢将种子贮藏在灌丛下方,但对埋藏基质没有明显偏好;2)埋藏微生境和埋藏基质对岩松鼠在对人工埋藏种子的找回过程中没有明显影响;3)岩松鼠对人工埋藏的核桃种子在种子找回过程中与自身埋藏的核桃种子有显著差异(Z=-2.499,P=0.014),对人工埋藏的胡桃楸种子在找回过程中与自身埋藏的胡桃楸种子有极显著差异(Z=-6.557,P<0.001)。     

啮齿动物对不同林木种子的搬运和取食微生境选择机制

不同植物种子依靠不同的方式实现扩散,啮齿动物对林木种子搬运后在取食点微生境和贮藏方式的选择存在偏好,研究其贮藏行为与微生境的关系是探究幼苗建成的关键。在秦岭中段火地塘林区,采用标签标记法,以锐齿槲栎、华山松和油松种子为材料,探究了小型啮齿动物对松栎混交林建群种种子扩散过程的影响。结果表明:1)油松种子原地取食率显著高于锐齿槲栎和华山松种子,且啮齿动物更倾向于搬运后取食(60%)和埋藏(4.33%)华山松种子,搬运后取食距离也为华山松最大(2.49 m);锐齿槲栎小种子被搬运后埋藏的距离最大(4.92 m)。2)除华山松种子外,其他类型种子被搬运后单个取食的比例均在85%以上;油松种子不存在埋藏点,而其他类型种子90%以上均以单个形式被埋藏。3)大部分种子被啮齿动物搬运后选择在裸地丢弃;锐齿槲栎大种子(87.5%)、小种子(78.57%)和华山松种子(53.33%)较大比例被啮齿动物埋藏在灌丛下方,埋藏在裸地的种子较少。4)大部分种子在灌丛下方被取食,仅华山松种子被啮齿动物搬运到洞穴取食;除油松种子被大量原地取食外,其他类型种子被搬运到取食点的种子比例基本呈现由微生境植被复杂到简单(灌丛—草丛—灌丛边缘—裸地)而逐渐减小的趋势。种子的营养价值及取食和搬运过程中啮齿动物付出的成本是影响种子命运的关键性因子,且啮齿动物对种子埋藏和取食地点的微生境存在较明显的选择性。     

岩松鼠的食物贮藏行为

食物贮藏是许多动物对不可预见的食物供应变化的一种重要适应.岩松鼠(Sciurotamias davidianus)为中国特有物种,广泛分布于华北地区的山地和丘陵地带.作者在北京市东灵山地区建造半自然围栏(4 m×3 m× 1 m),以核桃(Juglans regia)和山杏(Prunus armeniaca)种子为备选食物,对岩松鼠(12只)的食物贮藏行为进行了研究.数据的统计分析采用SPSS for Windows进行.研究结果表明1)岩松鼠表现出集中和分散两种食物贮藏方式,而分散贮藏是其偏好的贮藏方式;2) 当遇到贮藏食物被盗窃时,岩松鼠倾向于搬运更多的食物进行集中和分散贮藏;3)岩松鼠没有在食盘就地取食任何一种食物;把食物搬离后,岩松鼠所取食的山杏种子数量明显多于所取食的核桃数量;4)岩松鼠只选择核桃进行分散贮藏;5)岩松鼠对核桃的搬运距离大于山杏种子.本研究结果提示,在自然条件下,岩松鼠对核桃和山杏的天然更新起着不同的作用[动物学报 51(3)376-382,2005].     

贮藏种子的啮齿动物对油茶种子沉积形式的影响

种子沉积的质量常常涉及种子扩散作用者(如鸟类和小型啮齿动物)对生境和微生境的选择以及种子沉积的形式(如种子埋藏)。然而,很少的研究涉及到种子在离开母树后被这些动物沉积在何处。在四川省都江堰一个实验林场的2个林分(次生林和原生林)内,通过追踪用带编号的金属薄片标记的油茶(CamelliaoleiferaAbel.)种子的命运,研究了贮藏种子的啮齿动物对种子沉积的影响。研究发现在2个林分内,80%以上的种子被很好地埋藏在0~60mm深的土壤中,而小部分种子则被放置在地表(但有少量落叶遮盖)。小型啮齿动物喜好在灌丛下或灌丛边缘贮藏和取食种子,可能是在这样的微生境下它们在觅食时将遭遇较小的捕食风险。研究还发现,贮藏点的微生境分布随贮藏点等级而逐渐变化在两个林分内较高等级的贮藏点(如次贮藏点和三级贮藏点)比初级贮藏点有更多的种子被贮藏在灌丛下或灌丛边缘。这表明,啮齿动物对油茶种子的埋藏可能更有益于种子的存活、萌发以及幼苗的建成。啮齿动物将散落在母树下或其附近的油茶种子扩散到不同的微生境,这可能有利于他们遭遇到更多的适宜环境而萌发,实现幼苗补充。     

贮藏种子的啮齿动物对油茶种子沉积形式的影响

种子沉积的质量常常涉及种子扩散作用者(如鸟类和小型啮齿动物)对生境和微生境的选择以及种子沉积的形式(如种子埋藏).然而,很少的研究涉及到种子在离开母树后被这些动物沉积在何处.在四川省都江堰一个实验林场的2个林分(次生林和原生林)内,通过追踪用带编号的金属薄片标记的油茶(Camellia oleifera Abel.)种子的命运,研究了贮藏种子的啮齿动物对种子沉积的影响.研究发现:在2个林分内,80%以上的种子被很好地埋藏在0～60 mm深的土壤中,而小部分种子则被放置在地表(但有少量落叶遮盖).小型啮齿动物喜好在灌丛下或灌丛边缘贮藏和取食种子,可能是在这样的微生境下它们在觅食时将遭遇较小的捕食风险.研究还发现,贮藏点的微生境分布随贮藏点等级而逐渐变化:在两个林分内较高等级的贮藏点(如次贮藏点和三级贮藏点)比初级贮藏点有更多的种子被贮藏在灌丛下或灌丛边缘.这表明,啮齿动物对油茶种子的埋藏可能更有益于种子的存活、萌发以及幼苗的建成.啮齿动物将散落在母树下或其附近的油茶种子扩散到不同的微生境,这可能有利于他们遭遇到更多的适宜环境而萌发,实现幼苗补充.     

灌丛高度对啮齿动物贮藏和扩散辽东栎坚果的影响

本文选择高、矮两类灌丛生境, 于2002 年至2003 年在其中释放标记的辽东栎坚果, 并连续记录释放坚果的命运, 以了解生境差异对啮齿动物搬运、取食和贮藏辽东栎坚果的影响以及啮齿动物对微生境的选择等。研究结果表明, 大林姬鼠等啮齿动物对辽东栎坚果的短期取食非常强烈, 而分散贮藏的量则相对较少; 啮齿动物对辽东栎坚果的搬运距离在矮灌丛生境中显著大于高灌丛生境; 啮齿动物倾向于选择灌丛边缘和灌丛下方取食或贮藏辽东栎坚果, 这类微生境也有利于埋藏坚果的萌发。     

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。     

橡胶树在西双版纳引种区的种子捕食与散布

在纯橡胶林、胶-竹复合林、沟谷季节雨林、次生林及毗邻次生林的橡胶林共5种不同生境中释放标记的橡胶树种子,观测种子捕食者以及种子被捕食、埋藏和扩散情况。结果表明:纯橡胶林和沟谷季节雨林内的橡胶树种子大多数都原地存留(80%和98%),而胶-竹复合林内几乎100%的种子均被捕食;次生林内释放的橡胶树种子捕食率也几为100%,而毗邻次生林的橡胶林种子捕食率约50%;卡方检验表明,胶-竹复合林的种子捕食率显著高于沟谷季节雨林和纯橡胶林,次生林内显著高于毗邻的橡胶林(P0.001)。绝大多数橡胶树种子被捕食者原地或搬运后取食,仅有少量种子被埋藏。大多数种子(80%)被搬运距离在10 m以内。红外感应相机拍摄和饲喂实验发现,黄胸鼠(Rattus tanezumi)和红刺鼠(Rattus surifer)为主要的种子捕食者。可见,在西双版纳引种区橡胶树种子主要是被小型鼠类取食,但捕食强度较大,而被埋藏种子比例很低,散布距离较短(10 m)。     

贮藏点深度、大小及基质含水量对花鼠找寻红松种子的影响

2012年8-10月,在黑龙江省伊春市带岭林业局东方红林场,通过人工围栏控制实验,研究了贮藏点深度、大小及基质含水量对花鼠找寻红松种子的影响。发现：(1) 埋藏深度显著影响花鼠对贮藏点的找寻率,埋藏深度为 1 cm 和 2 cm 的找寻率显著高于 4 cm 和 6 cm。(2) 埋藏点大小对花鼠发现种子也有显著影响,埋藏点越大,花鼠发现贮藏点的比例越高。(3) 高的基质含水量利于花鼠找寻贮藏点。结果表明,围栏条件下食物贮藏点深度、大小及基质含水量的改变能显著影响花鼠对贮藏点的找寻。     

同种竞争压力对小泡巨鼠贮藏油茶种子行为的作用分析

2002年11～12月,在四川省都江堰地区的亚热带常绿阔叶林内利用人工修建的半自然状态围栏进行实验,研究了小泡巨鼠在有同种竞争存在条件下对油茶种子的埋藏行为。结果表明,小泡巨鼠在有竞争存在条件下,显增加了埋藏油茶种子的量。这一结果支持了“竞争的存在刺激鼠类埋藏”的假说。同时,研究结果表明,小泡巨鼠在有竞争存在条件下,显增加了对埋藏种子的搬运距离,每个贮藏点埋藏种子的数量也有所增加,同时埋藏的生境更多地偏向于遮蔽较好的微生境(草丛底层、灌丛下层)中。这些行为策略有可能有利于种子被埋藏植物的种群扩散。在讨论中,我们还通过比较鸟类和兽类在感觉器官上的差别,分析它们在有竞争存在条件下所采取的不同贮食策略。     

围栏条件下社鼠的食物贮藏行为

在北京东灵山地区, 用活捕笼从野外捕获社鼠, 并在野外建造围栏, 以山杏种子和辽东栎坚果为备选食物, 对围栏条件下社鼠的食物贮藏行为进行了研究。数据的统计分析采用SPSS for Windows (version 1010) 进行。结果表明: 1) 社鼠主要表现出食物的集中贮藏方式; 2) 贮藏食物被模拟盗窃使社鼠的食物贮藏活动加强; 3)社鼠倾向于选择山杏种子进行贮藏, 而对辽东栎坚果则取食较多。文中还讨论了食物类型、模拟盗窃等因素对社鼠食物贮藏行为的影响。     

Can forest fragmentation disrupt a conditional mutualism? A case from central Amazon

This is the first study to investigate whether scatter-hoarding behavior, a conditional mutualism, can be disrupted by forest fragmentation. We examined whether acouchies (Myoprocta acouchy, Rodentia) and agoutis (Dasyprocta leporina, Rodentia) changed scatter-hoarding behavior toward seeds of Astrocaryum aculeatum (Arecaceae) as a consequence of a decrease in forest-patch area. Our study was conducted at the 30-year-old Biological Dynamics of Forest Fragments Project, in central Amazon, Brazil. We tested whether forest size affected the number of Astrocaryum seeds removed and scatter-hoarded (and likely dispersed) by acouchies and agoutis, as well as the distance that the seeds were hoarded. The study extended over three seasons: the peak of the rainy season (March–April), the transition between the rainy and the dry season (May–June), and the peak of the dry season (August–September). Our results revealed that the number of seeds removed was larger in smaller fragments, but that the percentage of seeds hoarded was much lower, and seeds eaten much higher, in 1-ha fragments. Moreover, fewer seeds were taken longer distances in fragments than in the continuous forest. Site affected the number of seeds removed and season affected the percentage of seeds hoarded: more seeds were removed from stations in one site than in two others, and hoarding was more important in April and September than in June. Our study reveals that scatter-hoarding behavior is affected by forest fragmentation, with the most important disruption in very small fragments. Fragmentation converts a largely mutualistic relationship between the rodents and this palm in large forest patches into seed predation in small fragments.     

Effects of habitat and season on removal and hoarding of seeds of wild apricot (Prunus armeniaca) by small rodents

The wild apricot (Prunus armeniaca) is widely distributed in the Donglingshan Mountains of Mentougou District of Beijing, China, where its seeds may be an important food resource for rodents. Predation, removal and hoarding of seeds by rodents will inevitably affect the spatio-temporal pattern of seed fate of wild apricot in this area. By marking and releasing tagged seeds of wild apricot, we investigated seeds survival, scatter-hoarding, cache size and seedling establishment, and the preference of micro-habitats used by rodents to store seeds. The results showed that: (1) rodents in this area hoarded food intensively in autumn, as well as in spring and summer. (2) There were significant effects of habitat and season on removal rate of tagged seeds at releasing plots. In both two types of habitats, Low and High shrub, tagged seeds were removed most rapidly by rodents in autumn, at intermediate rates in spring and least rapidly in summer. (3) During three seasons, mean dispersal distance of scatter-hoarded seeds in Low shrub habitat was greater than that in High shrub. Most removed seeds were buried within 21.0 m of the releasing plots. (4) In both two types of habitats, Low and High shrub, rodents tended to carry seeds to US (Under shrub) and SE (Shrub edge) microhabitats for scatter-hoarding or predation. (5) Among the caches made by rodents, most caches contained only one seed, but up to three seeds were observed; caches of 2–3 seeds were common in autumn. (6) By comparing dental marks, we determined that large field mice (Apodemus peninsulae) and David’s rock squirrels (Sciurotamias davidianus) contributed to removal and predation of released tagged seeds. However, only the large field mice exerted a pivotal and positive role on the burial of dispersed seeds. (7) Establishment of three seedlings originated from seeds buried by rodents was documented in High shrub habitat.     

Effects of burrow condition and seed handling time on hoarding strategies of Edward's long-tailed rat (Leopoldamys edwardsi)

Many hoarding rodents use burrows not only for dwelling and protection from natural enemies, but also for food storage. However, little is known how burrows used by scatter-hoarding animals influence their foraging behaviors. In addition, handling time for a given food item has a fundamental impact on hoarding strategies of these hoarding animals: food items with longer handling time are more likely to be hoarded due to increasing predation risk because the animals spend more time outside their burrows if they consumed such food. By providing with two types of artificial burrows (aboveground vs. underground) and two types of food items (i.e. seeds) with contrasting handling times, we investigated how burrow condition and handling time co-influence hoarding strategies of a key scatter-hoarding rodent, Edward's long-tailed rat (Leopoldamys edwardsi) in large enclosures in southwest China. We found that only a few animals larder-hoarded fewer seeds when only aboveground burrows were available, while over 80% of the animals preferred to use the underground burrows and hoard significantly more seeds in the burrows when both aboveground and underground burrows were provided simultaneously. We also found that seed handling time significantly affected hoarding strategies of the animals: they consumed and/or scatter-hoarded more Camellia oleifera seeds with shorter handling time outside the burrow, but consumed and larder-hoarded more Lithocarpus harlandii seeds with longer handling time in underground burrows. Our study indicates that both burrow types and seed handling time have important impacts on hoarding strategies of scatter-hoarding animals.     

Sensitivity to Seed Germination Schedule by Scatter‐Hoarding Pére David's Rock Squirrels During Mast and Non‐Mast Years

Mast seeding by oaks (Quercus) and other seed species has fundamental impacts on the behavior of individual scatter‐hoarding animals as well as their population and community dynamics. Scatter‐hoarding squirrels are highly sensitive to acorn germination schedule and stop germination by removing the embryo of non‐dormant acorns to prevent losses of energy and nutrients by rapid germination. However, we know little about how this behavior is affected by mast seeding. We investigated foraging decisions made by free‐ranging Pére David's rock squirrel (Sciurotamias davidianus) with three species that produce non‐dormant seeds and two species that produce dormant seeds (Fagaceae) during mast and non‐mast years in Central China from 2007–2010. Consistent with the food perishability hypothesis that squirrels make decisions to minimize the perishability of their caches, non‐dormant seeds were hoarded at a lower rate, but had a higher probability of embryo removal than dormant seeds. Compared with non‐mast years, a lower proportion of seeds were harvested during mast years, but a higher proportion of the harvested seeds were hoarded. In addition, the probability of embryo removal in the hoarded seeds was higher during mast years. Squirrels also dispersed seeds at shorter distances during mast years. Moreover, the interactions between mast seeding and seed germination schedule had significant effects on squirrel foraging decisions, including embryo removal. This study is the first to demonstrate that squirrels show greater sensitivity to seed germination potential when seeds are abundant.     

Predation and dispersal of large and small seeds of a tropical palm

Seed size may vary greatly among individuals within plant species. What effects the extremes of this variation have for seeds taken by small mammals are poorly understood. Not all seeds removed by small mammals are necessarily eaten. Small rodents are common seed predators, but they may disperse a significant proportion of seeds by scatter hoarding them via burial. Size-dependent predation and dispersal of seeds has not been directly tested within a plant species for tropical rodents. This study tested whether or not large and small nuts of Astrocaryum mexicanum (Palmae) differed in their fates due to handling by the spiny pocket mouse Heteromys desmarestianus (Heteromyidae). Exclosures were used to give small rodents exclusive access to A. mexicanum nuts. H. desmarestianus preferentially consumed large over small A. mexicanum nuts, but cached (in burrows and by scatter hoarding) similar proportions of these nuts by size. Small nuts tended to be buried farther away from exclosures than large nuts. Although sample sizes of buried nuts were small, the rodents retrieved all buried large nuts, but 30% of the small nuts remained buried long enough to germinate. I also examined predispersal predation by insects and found that insects appear to have no size preference for A. mexicanum nuts, but insect predation appears to hinder nut development. Thus, nuts attacked by insects develop to be significantly smaller, with a low proportion of undamaged endosperm, than uninfested nuts. It is hypothesized that the preferential predation of large A. mexicanum nuts by H. desmarestianus is a response by these rodents to insect predation.     

围栏条件下中华姬鼠和北社鼠对完好和虫蛀锐齿槲栎种子的取食和贮藏行为差异

正种子贮藏是啮齿动物利用食物资源的一种适应性行为(Vander Wall,1990)。经过长期的进化,啮齿动物形成了两种主要的贮藏食物行为:一是集中贮藏,即把收集的大量食物集中在其洞穴或临时栖息场所贮藏,通常有少量贮藏点,每个贮藏点有大量种子;二是分散贮藏,即把收集的种子分散贮藏在相对较大范围或其巢域周围,通常有大量贮藏点,但每个贮藏点仅贮藏约1～3粒种子。与集中贮藏相比,分散贮藏扩展了植物种子的分布范围,降低了种子被动物取食的概率。由于被分散贮藏的     

埋藏点深度、间距及大小对花鼠发现向日葵种子的影响

2005 年5 ～6 月,在北京东灵山地区,通过在实验围栏内设置不同深度、间距和大小的埋藏点,研究了花鼠对向日葵种子的发现率。结果表明,花鼠发现埋藏种子的比例随埋藏深度的增加而减少,埋藏深度> 5 cm后,发现比例较低,且不再有明显变化;花鼠发现埋藏种子的比例随埋藏点间距的减小而增加,埋藏点间距<0.5 m后,发现比例较高,且不再有明显变化;花鼠发现埋藏种子的比例随着埋藏点增大而增加,埋藏点种子> 5 g后,发现比例较高,且不再有明显变化。     

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)。