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。     

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     

华山松种子大小对啮齿动物贮藏行为的影响及其时空变化

华山松(Pinus armandii)是广泛分布在中国中西部海拔1000～3300m的山地、种子较大(约300mg)的松属植物。为探究华山松种子大小对啮齿动物贮藏行为的影响,于2006年和2007年在滇西北3个不同地点进行种子标记和追踪实验。结果表明,在所有年份和地点,啮齿动物都倾向于贮藏更多的大种子和取食更多的小种子;啮齿动物贮藏大种子的数量,以及平均距离和最大距离均显著高于小种子。3个地点具有不同的啮齿动物群落结构,从而对种子命运产生显著影响。种子命运在两个年份间也存在显著差异。     

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。     

种子萌发过程中主要贮藏物质的转变实验

种子萌发过程中主要贮藏物质的转变实验陈平平（南京教育学院生物学系２１００１７）孙美姝（南京市第三中学２１０００２）一般植物种子中贮藏的大量有机物,主要是淀粉、脂肪和蛋白质。不同植物的种子这三类物质的含量,有着很大的差异。例如绿豆含有丰富的淀粉,约为４...     

啮齿动物分散贮食的影响因素

分散贮藏食物是许多啮齿动物所采取的一种重要的觅食策略。分散贮藏方式对于植物的幼苗建成、更新、群落演替和生态系统功能的维持有着积极的意义。多种因素影响啮齿动物对食物分散贮藏。内部因素有性别、年龄、社群关系、饥饿状态等;外部因素包括食物特征、竞争与盗食环境条件等。这些因素并非独立地发挥作用,内部因素受外部刺激的综合影响,而且外部因素只能通过改变动物的内部状态才能发挥作用。特别是在一些室内的或围栏实验中,实验设计的差异不仅影响动物的食物贮藏及其管理,也会造成实验结果的不同。     

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。     

围栏条件下同域分布三种鼠对两种种子的贮藏行为差异

种子贮藏是啮齿动物利用食物资源的一种适应性行为。同域分布的啮齿动物可能进化出不同的贮藏方式和策略。为了解同域分布的啮齿动物的贮藏策略及种子特征在动物贮藏行为决策中的作用,于2014年10-12月,通过半自然围栏实验,在秦岭南坡的佛坪国家级自然保护区内调查了同域分布社鼠、中华姬鼠和甘肃仓鼠对板栗和锐齿槲栎种子的贮藏策略差异,并探讨了2种种子特征在鼠类贮藏行为决策中的作用。结果显示：1）社鼠主要集中贮藏板栗种子,未分散贮藏任何种子;中华姬鼠未贮藏板栗种子,对锐齿槲栎种子同时表现出集中和分散贮藏行为,以集中贮藏为主;甘肃仓鼠对2种种子均表现出集中贮藏行为,未表现分散贮藏行为。2）社鼠和甘肃仓鼠均倾向于集中贮藏有较高营养价值（高蛋白和脂肪）且单宁含量较低的板栗种子;中华姬鼠倾向于分散贮藏单宁含量高的锐齿槲栎种子。3）3种鼠均喜好取食有较高营养价值且单宁含量低的板栗种子,很少取食单宁含量高的锐齿槲栎种子。结果表明同域分布鼠类对不同种子的贮藏方式有所不同,种子特征影响鼠类的取食和贮藏策略。     

啮齿动物对印度栲种子扩散的效率

从2007 年11 月到2009 年11 月,在西双版纳热带雨林中选取三棵母树,在每棵树下每年释放标记印度栲种子200 粒(三年共计1 800 粒),并追踪其命运。通过调查啮齿动物搬运和分散贮藏印度栲种子的比例,以及
查贮藏种子的微生境、贮藏点大小和扩散距离,分析贮藏种子的存活情况,进而评估啮齿动物对印度栲种子扩散的效率。结果表明,啮齿动物搬运了69.3% 的印度栲种子,被搬运的种子中18% 被分散贮藏。所有被分散贮藏的种子均被埋于落叶下或埋于土壤表层,并且大部分贮藏点仅含一粒种子。种子的扩散距离从0.5 m 到43.8 m,平均距离为7.1 m,扩散距离在年间没有显著差异。2007 年(种子密度低,啮齿动物密度高) 没有种能最终存活到实验结束,而2008 和2009 年(种子密度高,啮齿动物密度低)分别有0.3% 和1.5% 的种子存活。研究表明,啮齿动物是印度栲有效的种子扩散者,但其扩散效率很大程度上取决于森林中种子的密度和啮齿动物的丰富度。     

Dispersal by rodent caching increases seed survival in multiple ways in canopy‐fire ecosystems

Seed‐caching rodents have long been seen as important actors in dispersal ecology. Here, we focus on the interactions with plants in a fire‐disturbance community, specifically Arctostaphylos species (Ericaceae) in California chaparral. Although mutualistic relationships between caching rodents and plants are well studied, little is known how this type of relationship functions in a disturbance‐driven system, and more specifically to systems shaped by fire disturbance. By burying seeds in the soil, rodents inadvertently improve the probability of seed surviving high temperatures produced by fire. We test two aspects of vertical dispersal, depth of seed and multiple seeds in caches as two important dimensions of rodent‐caching behavior. We used a laboratory experimental approach to test seed survival under different heating conditions and seed bank structures. Creating a synthetic soil seed bank and synthetic fire/heating in the laboratory allowed us to have control over surface heating, depth of seed in the soil, and seed cache size. We compared the viability of Arctostaphylos viscida seeds from different treatment groups determined by these factors and found that, as expected, seeds slightly deeper in the soil had substantial increased chances of survival during a heating event. A key result was that some seeds within a cache in shallow soil could survive fire even at a depth with a killing heat pulse compared to isolated seeds; temperature measurements indicated lower temperatures immediately below caches compared to the same depth in adjacent soil. These results suggest seed caching by rodents increases seed survival during fire events in two ways, that caches disrupt heat flow or that caches are buried below the heat pulse kill zone. The context of natural disturbance drives the significance of this mutualism and further expands theory regarding mutualisms into the domain of disturbance‐driven systems.     

啮齿类对植物种子的传播作用

种子植物是固着生活的有机体 ,如果能成功地将种子扩散到适宜的生境 ,将会在生存竞争中获得优势。在长期的进化过程中 ,不同的植物依赖不同的媒介传播种子 ,如风传播 ,水传播 ,或自身的力量传播 (重力、弹爆力等 )。在很多情况下也依靠动物完成种子扩散 ,即所谓的动物传播。根据依赖的动物对象不同可以分为蚁传播 ,鸟传播 ,哺乳类传播 ,以及鱼传播 ,爬行类传播等。哺乳类中传播种子的类群主要包括翼手类 ,灵长类和啮齿类。由于能够飞行 ,热带食果实的蝙蝠(属翼手类 )对种子的传播作用最明显 ,研究得也最多 ,源于蝙蝠类的传播特称为ｃｈｉｒ…     

Two-phase seed dispersal: linking the effects of frugivorous birds and seed-caching rodents

Frugivorous birds disperse the seeds of many fruit-bearing plants, but the fate of seeds after defecation or regurgitation is often unknown. Some rodents gather and scatter hoard seeds, and some of these may be overlooked, germinate, and establish plants. We show that these two disparate modes of seed dispersal are linked in some plants. Rodents removed large (>25 mg) seeds from simulated bird feces (pseudofeces) at rates of 8–50%/day and scatter hoarded them in soil. Ants (Formica sibylla) also harvested some seeds and carried them to their nests. Rodents carried seeds 2.5±3.2 m to cache sites (maximum 12 m) and buried seeds at 8±7 mm depth. Enclosure studies suggest that yellow pine chipmunks (Tamias amoenus) and deer mice (Peromyscus maniculatus) made the caches. In spring, some seeds germinated from rodent caches and established seedlings, but no seedlings established directly from pseudofeces. This form of two-phase seed dispersal is important because each phase offers different benefits to plants. Frugivory by birds permits relatively long-range dispersal and potential colonization of new sites, whereas rodent caching moves seeds from exposed, low-quality sites (bird feces on the ground surface) to a soil environment that may help maintain seed viability and promote successful seedling establishment.     

Seed dispersal of Korean pine Pinus koraiensis labeled by two different tags in a northern temperate forest, northeast China

Two types of tagging methods, i.e., a 1 × 3-cm tin tag attached to seed with a 10- to 12-cm metal wire (total weight 0.32 g) and a 2 × 4-cm white plastic tag fastened to seed with an identical metal wire (total weight 0.57 g) were used to study their effects on seed dispersal of Korean pine by small rodents. A total of 600 seeds were released to assess four main points: (1) difference in seed survival rates, (2) difference in caching behaviors of small rodents, (3) difference in dispersal distances, and (4) proportion of seed missing. The results demonstrated that seed removal for wire-plastic-tagged seeds was faster than that for wire-tin-tagged seeds. There was no apparent difference in the proportion of seeds eaten in situ (42% and 52% for wire-plastic-tagged seeds and wire-tin-tagged seeds, respectively). We found 41% and 1% of seeds were moved and hoarded for wire-plastic-tagged seeds and wire-tin-tagged seeds, respectively. However, 2.33% and 14% of seeds were missing, and their ultimate fates were not known for wire-plastic-tagged seeds and wire-tin-tagged seeds, respectively. We found the wire-plastic-tagged seeds easier to track than the wire-tin-tagged seeds due to the fact that the white plastic tags were more salient than the tin tags in field environments. The average dispersal distances were 4.11 ± 2.40 m and 3.01 ± 2.06 m for wire-plastic-tagged seeds and wire-tin-tagged seeds, respectively, and showed great difference. Despite most being eaten in situ or after removal, 41% of seeds were cached for wire-plastic-tagged seeds, much more than for wire-tin-tagged seeds. A total of 71 primary caches (123 seeds) were found for wire-plastic-tagged seeds, with the average and maximum cache sizes being 1.73 and 6, respectively. However, only three caches were found, and cache size was equal to one for wire-tin-tagged seeds. The above data suggests there is some uncertainty in different tagging methods to used track seed fates. Despite their effectiveness in helping to trace seed dispersal or movement by seed-dispersing rodents, different tagging methods—including size, color, and mass—need to be fully understand in enclosure experiments .     

Re-caching of acorns by rodents: Cache management in eastern deciduous forests of North America

Scatter-hoarding rodents such as tree squirrels selectively cache seeds for subsequent use in widely-spaced caches placed below the ground surface. This behavior has important implications for seed dispersal, seedling establishment, and tree regeneration. Hoarders manage these caches by recovering and eating some seeds, and moving and re-caching others. This process of re-caching, however, is poorly understood. Here, we use radio-telemetry to evaluate re-caching behavior for the management of acorn caches by rodents in eastern deciduous forests. We also test the hypothesis that as seeds are re-cached, the distance from the source increases. Radio transmitters were implanted in Northern red oak (Quercus rubra) acorns and presented to rodents in a natural setting over 3 seasons. We used radio-telemetry to track and document evidence of recovery and re-caching. We tracked a total of 102 acorns. Of the 39 radio-tagged acorns initially cached, 19 (49%) were cached on two or more occasions; one acorn was cached four times. The hypothesis that rodents move seeds to progressively greater distances from the source is not well-supported, suggesting that acorns are being moved within an individual's home range. Given the species of rodents in the study area, gray squirrels (Sciurus carolinensis) are the most likely to be responsible for the caching and re-caching events. Gray squirrels appear to engage in extensive re-caching during periods of long-term food storage, which has important implications for understanding how caching behavior influences acorn dispersal and oak regeneration.     

Ecosystem Services from Keystone Species: Diversionary Seeding and Seed‐Caching Desert Rodents Can Enhance Indian Ricegrass Seedling Establishment

Seeds of Indian ricegrass (Achnatherum hymenoides), a native bunchgrass common to sandy soils on arid western rangelands, are naturally dispersed by seed‐caching rodent species, particularly Dipodomys spp. (kangaroo rats). These animals cache large quantities of seeds when mature seeds are available on or beneath plants and recover most of their caches for consumption during the remainder of the year. Unrecovered seeds in caches account for the vast majority of Indian ricegrass seedling recruitment. We applied three different densities of white millet (Panicum miliaceum) seeds as “diversionary foods” to plots at three Great Basin study sites in an attempt to reduce rodents' over‐winter cache recovery so that more Indian ricegrass seeds would remain in soil seedbanks and potentially establish new seedlings. One year after diversionary seed application, a moderate level of Indian ricegrass seedling recruitment occurred at two of our study sites in western Nevada, although there was no recruitment at the third site in eastern California. At both Nevada sites, the number of Indian ricegrass seedlings sampled along transects was significantly greater on all plots treated with diversionary seeds than on non‐seeded control plots. However, the density of diversionary seeds applied to plots had a marginally non‐significant effect on seedling recruitment, and it was not correlated with recruitment patterns among plots. Results suggest that application of a diversionary seed type that is preferred by seed‐caching rodents provides a promising passive restoration strategy for target plant species that are dispersed by these rodents.     

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。     

Recaching Decisions of Florida Scrub‐Jays are Sensitive to Ecological Conditions

Food caching animals depend on their caches at times of low food availability. Because stored food is susceptible to being stolen or degraded, many species employ cache protection strategies such as ceasing caching in the presence of others or avoiding storing perishable items for long periods. Several species frequently recover their caches and recache, which may reduce pilferage or degradation of cached items. We studied the food handling decisions of Florida scrub‐jays (Aphelocoma coerulescens) after cache recovery to determine the roles that social and ecological environments play in post‐recovery decisions. Instead of reducing recaching in the presence of others, recovering jays flew away from the recovery site, allowing them to eat or recache a recovered item regardless of the social context. Microhabitat type and soil moisture of the recovery sites had a significant influence on whether recoveries were eaten or recached; most items that were recached had been recovered from bare sand sites or sites with low soil moisture. Taken together, our results suggest that food store management of Florida scrub‐jays are unaffected by the social context, but are strongly affected by the habitat conditions that influence the quality of caches.     

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.     

Endocarp thickness affects seed removal speed by small rodents in a warm-temperate broad-leafed deciduous forest, China

Seed traits are important factors affecting seed predation by rodents and thereby the success of recruitment. Seeds of many tree species have hard hulls. These are thought to confer mechanical protection, but the effect of endocarp thickness on seed predation by rodents has not been well investigated. Wild apricot (Prunus armeniaca), wild peach (Amygdalus davidiana), cultivated walnut (Juglans regia), wild walnut (Juglans mandshurica Maxim) and Liaodong oak (Quercus liaotungensis) are very common tree species in northwestern Beijing city, China. Their seeds vary greatly in size, endocarp thickness, caloric value and tannin content. This paper aims to study the effects of seed traits on seed removal speed of these five tree species by small rodents in a temperate deciduous forest, with emphasis on the effect of endocarp thickness. The results indicated that speed of removal of seeds released at stations in the field decreased significantly with increasing endocarp thickness. We found no significant correlations between seed removal speed and other seed traits such as seed size, caloric value and tannin content. In seed selection experiments in small cages, Père David's rock squirrel (Sciurotamias davidianus), a large-bodied, strong-jawed rodent, selected all of the five seed species, and the selection order among the five seed species was determined by endocarp thickness and the ratio of endocarp mass/seed mass. In contrast, the Korean field mouse (Apodemus peninsulae) and Chinese white-bellied rat (Niviventer confucianus), with relatively small bodies and weak jaws, preferred to select small seeds like acorns of Q. liaotungensis and seeds of P. armeniaca, indicating that rodent body size is also an important factor affecting food selection based on seed size. These results suggest endocarp thickness significantly reduces seed removal speed by rodents and then negatively affects dispersal fitness of seeds before seed removal of tree species in the study region. However, effect of endocarp thickness on final dispersal fitness needs further investigation because it may increase seed caching and survival after seed removal.