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。     

四种昼行性动物取食和贮藏红松种子的行为比较

于2005年10月1~7日,采用全时焦点观察法研究了黑龙江凉水国家级自然保护区19林班原始红松林内松鼠(Sciurus vulgaris)、星鸦(Nucifraga caryocatactes)、花鼠(Eutamias sibiricus)和普通(Sitta europaea)4种昼行性动物取食和贮藏红松种子的行为。结果表明①4种动物的取食和贮藏过程包括找寻、松塔的处理与种子剥食、松塔和种子的搬运、确定贮藏点、掩藏和警戒等6个环节,但它们在6个环节所耗费的时间均存在较大差异;②4种动物的种子消耗量亦存在较大差异(χ2=144.67,df=3,P<0.05),松鼠、星鸦、花鼠和普通每次平均取食红松种子的数量分别为40.0粒、20.9粒、16.7粒和1.3粒。松鼠与星鸦能够在地面和树冠层取食或收获红松松塔及种子,而花鼠与普通则主要在地面取食;③4种动物贮藏种子的能力亦有较大差异(χ2=68.76,df=3,P<0.05Kruskal-Wallis),松鼠、星鸦和普通主要分散贮藏红松种子,松鼠和星鸦的贮藏量最多,而花鼠很少分散贮藏种子,但能将许多红松种子集中贮藏在少数的贮藏点;④取食松籽的种间竞争强烈,在红松林的垂直高度上,松鼠和星鸦具有高度重叠的取食生态位,花鼠和普通因为种群数量大,对地面种子库的消耗量较大。分析结果显示,松鼠与星鸦是红松种子扩散的传播者,普通是种子消耗者兼传播者,而花鼠则近乎是完全的捕食者。     

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。     

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%.     

凉水自然保护区松鼠与星鸦对红松种子分散贮藏的特征分析

2005年9月30日～10月8日在凉水保护区,通过野外分层抽样和系统抽样调查松鼠(Sciurus vulgaris)和星鸦(Nucifraga caryocatactes)的贮点特征,初步估算了主要生境内的贮食密度和贮藏量。松鼠贮点的平均大小为(3·02±0·11)粒,平均深度为(2·67±0·08)cm;星鸦贮点的平均大小为(3·21±0·22)粒,平均深度为(2·73±0·15)cm。松鼠、星鸦的贮点大小和深度均无显著差异。原始红松林内的贮食密度和贮食量最大,这和松鼠的贮食行为密切相关。保护区内贮食动物的贮食总量为(11876757±812304)粒,其中松鼠的贮食量为(8686312±683360)粒,星鸦的贮食量为(3190444±410377)粒。动物的贮食行为是影响红松天然更新的重要因子之一,对红松的天然更新有重要意义。     

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)。     

植物大年结实及其与动物贮食行为之间的关系

大年结实(mast seeding)是多生年植物种群周期性同步大量繁殖的一种自然现象。大年结实作为植物适应环境条件、提高繁殖能力的一种策略而备受关注, 但其驱动机制和进化意义尚存在较大争议。在依赖动物扩散种子的植物中, 大年结实被认为是一种调控动物贮食行为、提高种子扩散效率, 并最终增加繁殖成功率的一种策略; 动物介导的植物间互作可能是促进植物共存的进化驱动力。本文简要梳理了大年结实现象的各种假说, 提出了一个包括气候、资源、动植物互作的理解大年结实机制的概念框架, 并着重讨论了大年结实和动物贮食行为之间的关系及其进化和生态意义。建议未来研究需要借助长期生态监测和分子生物学方法, 揭示植物大年结实与动物贮食行为之间的生态与进化过程。     

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．     

食物充足条件下花鼠贮藏行为的研究

为了研究花鼠(Eutamias sibiricus)在食物充足情况下是否对食物进行贮藏。采用围栏实验来观察探究花鼠对食物贮藏的行为,选用四只花鼠进行行为观察。在提供足量的毛榛和东北红松种子的情况下,观察每只花鼠的取食量、埋藏量、巢内贮藏量、搬运量、原地剩余量、贮藏点数量、搬运量等指标。实验表明在食物充足情况下,花鼠取食后仍然对多余的食物进行搬运和贮藏。     

种子特征和结实量对啮齿动物取食和扩散种子行为的影响

为了深入了解啮齿动物在不同种子丰富度条件下对不同大小和单宁含量种子的觅食行为策略及其与植物种群更新的关系,在宁夏六盘山区的华北落叶松人工林,研究了不同大小和单宁含量[0%Tannin(T)、2%T、8%T和15%T]的人工种子在模拟结实小年和结实大年对啮齿动物取食和扩散行为的影响.结果表明: 啮齿动物消耗种子速度在结实小年更快,结实大年的种子消耗速度相对缓慢. 种子就地取食率(ISPR)在不同结实年份间无显著差异,扩散后取食率(PRAD)在结实小年显著高于结实大年,但前者的扩散后贮藏率(HRAD)显著低于后者;种子扩散后的取食距离(PDAD)和贮藏距离(HDAD)在结实小年均显著大于结实大年.在结实小年,大种子的PDAD和HDAD均大于小种子,前者在不同大小种子间均差异显著,而后者仅在2%T和15%T的不同大小种子间差异显著;在结实大年,除0%T外的其他单宁含量种子的PDAD和HDAD在不同大小种子间均差异显著.ISPR在中等单宁含量种子最大,高单宁含量种子最小;PRAD分别在结实小年的高单宁含量种子和结实大年的无单宁种子最大;不论在结实大年还是结实小年,HRAD均在高单宁含量种子最大,中等单宁含量种子最小.这说明结实大年可延缓啮齿动物对种子的消耗速率,提高种子的HRAD,但种子扩散距离减小;啮齿动物在结实大年和小年均表现出对大种子的扩散偏好,且大种子被扩散的距离更远;啮齿动物在不同结实年份均偏好于就地取食中等单宁含量种子,而扩散高单宁含量种子.     

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.     

Effects of mast seeding and rodent abundance on seed predation and dispersal of <Emphasis Type="Italic">Quercus aliena</Emphasis> (Fagaceae) in Qinling Mountains,Central China

Rodents act as seed predators and dispersers and play an important role in the regeneration of plants. Seed production and rodent abundance may influence caching rodents’ decision to consume or cache seeds. We studied how did seed production and rodent abundance co-influence seed predation and dispersal by rodents in the mast seed years (2011 and 2013) and non-mast seed years (2012 and 2014) in the Qinling Mountains of Shaanxi Province, Central China. We found that: (1) The seed removal rates were much faster in the non-mast seed years than in the mast seed years. (2) Although the total number of seeds consumed (including eaten in situ and after removal) was higher in the non-mast seed years than in the mast seed years, no significant differences were found in the seeds that were eaten in situ and after removal among the 4 years. (3) Significant differences were observed in seeds that were cached among the 4 years, and more seeds were cached in the non-mast seed years than in the mast seed years. (4) The primary and secondary seed dispersal distances (including cached and eaten) were all longer in the non-mast seed years than in the mast seed years. Overall, these results indicate that non-mast seeding with low per capita seed availability could increase both seeds consumption and caching by rodents. Our results are partially supporting the predator satiation hypothesis.     

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。     

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.     

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.     

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.     

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.