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)。     

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。     

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。     

长白山针阔混交林不同演替阶段的昆虫多样性

昆虫多样性变化对生态系统健康有重要的指示作用, 为研究昆虫群落变化与生境演替之间的关系, 本研究采用网捕、灯诱和诱捕法系统调查了长白山针阔混交林不同演替阶段（次生白桦林、次生针阔混交林、原始阔叶红松林）昆虫群落的组成和多样性, 分析了昆虫在森林演替过程中的规律及与植被群落之间的关系。系统调查共采集昆虫标本8 183头, 隶属于14个目699种, 其中鳞翅目和鞘翅目是主要优势类群。次生针阔混交林昆虫的个体数量最多, 原始阔叶红松林中物种数最多。不同演替阶段昆虫群落的物种数和个体数差异不显著, 但次生针阔混交林、原始阔叶红松林的Fisher’s α指数显著高于次生白桦林。目水平上的昆虫多样性未表现出显著性差异. 昆虫多样性在森林演替过程中和草本植物多样性的变化趋势相同;由于食性和生境选择的不同, 森林演替过程中鳞翅目昆虫多样性逐渐升高, 而鞘翅目多样性逐渐降低。     

长白山地区不同林型红松种群生态位特征

采用Levins、Hurlbert生态位宽度和Pianka生态位重叠计测方法,分析长白山地区不同林型红松种群生态位特征。结果表明,红松种群是长白山地区顶极群落原始阔叶红松林的优势种群,其生态位宽度呈现原始阔叶红松林白桦林落叶松阔叶混交林。相比于其他种群的生态位宽度,总体上原始阔叶红松林中耐阴树种的生态位宽度较大,阳性树种的生态位宽度较小,而次生林则相反。在生态位重叠方面,红松与各林型中其他种群表现出不完全重叠。在原始阔叶红松林和白桦林中,红松与其他乔木种群对资源有明显的共享趋势。红松与原始阔叶红松林的色木槭、紫椴、青楷槭、白桦,与落叶松阔叶混交林的紫椴、蒙古栎、长白落叶松,与白桦林的色木槭、白桦,对同一资源有相同或相似的要求,且当资源不足时会产生竞争。     

1896年和1986年黑龙江省几种森林景观的特征变化

通过用地理信息系统软件对 1896和 1986年的黑龙江省的森林分布图中提取的 5种森林景观进行处理和计算得到 :在 90年中 ,云杉 (Picea)、冷杉 (Abies)林、樟子松 (Pinussylvestrisvar.mongolicaLitvin)林和阔叶红松 (PinuskoraieusisSieb .etZucc .)林斑块总面积分别减少 87%、40 %和 84%。北纬 5 1°以北地区阔叶林斑块面积增加 5 0 0 % ,北界向西北扩展约 2 90km。云杉、冷杉林和针阔混交林斑块在 1986年的斑块的分维数小于 1896年的斑块的分维数 ,而樟子松林、阔叶红松林和阔叶林的 1986年斑块分维数都分别大于 1896年的分维数。云杉、冷杉林、樟子松林、阔叶红松林和阔叶林景观破碎化增加。云杉、冷杉林 ,樟子松林和阔叶红松林转化为针阔混交林的比率大于各自保留的比率。     

不同类型人工阔叶红松林高等植物物种多样性

采用Shahnon—Wiener多样性指数、均匀度指数和优势度指数等指标研究了辽宁东部山区人工营造20年生5种阔叶树水曲柳、刺愀、紫椴、色赤汤、白桦与红松形成的混交林及人工红松纯林内各层次高等植物物种多佯性,并对该6种林分的垂直结构进行比较。结果表明,人工阔叶红松混交林内植物种类丰富,数量较多;红松纯林内的植物种数仅为人工阔叶红松混交林的42％～52％,植物总数量只有混交林的11％～37％。人工阔叶红松林各层次高等植物物种多样性均好于人工红松纯林,但与原始阔叶红松林相比。人工阔叶红松林的物种多样性还较低。在林分垂直结构方面,人工阔叶红松混交林的成层现象明显,层次结构复杂,垂直多样性丰富;红松纯林只有1个主林冠层,灌木层、草本层较矮,林分垂直结构单一。     

辽东山区人工阔叶红松林植物多样性与生产力研究

根据对人工营造20年生阔叶红松林典型样地的调查,分析了人工阔叶红松林的生产力、植物多样性及其相互关系.结果表明,人工营造的20年生红松-白桦、红松-色赤杨、红松-水曲柳混交林林地生产力分别达.529、4.9和5.82 t·hm^-2·yr^-1,高于同龄人工红松纯林(3.812 t·hm^-2·yr^-1);而红松-刺楸、红松-紫椴混交林分别为2.945和2.84 t·hm^-2·yr^-1,低于人工红松纯林.人工阔叶红松林乔木层、灌木层、草本层植物多样性均高于红松纯林;红松纯林内的植物种数仅为人工阔叶红松混交林的42%～52%,植物总数量也只有混交林的11%～37%.人工营造阔叶红松林是迅速恢复、发展顶极阔叶红松林的一种有效措施,但要经历相当长的时间和人为的不断调控.     

小兴安岭5种林型土壤呼吸时空变异

原始阔叶红松林、谷地云冷杉林、阔叶红松择伐林、次生白桦林、人工落叶松林是小兴安岭乃至东北地区的重要森林类型。采用红外气体分析法比较测定了这几种森林类型的土壤呼吸及其相关环境因子,分析探讨了这几种森林类型土壤呼吸的时空变异。结果表明:各林型土壤呼吸与5 cm深土壤温度(T5)呈显著的指数相关,并且土壤呼吸与土壤温度、土壤湿度及其相互作用的回归模型可以解释各林型土壤呼吸约71%的季节变异。生长季平均土壤呼吸速率为次生白桦林(3.59μmolCO.2m-.2s-1)>谷地云冷杉林(3.52μmolCO.2m-.2s-1)>阔叶红松择伐林(3.44μmolCO.2m-.2s-1)>原始阔叶红松林(2.58μmolCO.2m-.2s-1)>人工落叶松林(2.29μmolCO.2m-.2s-1),说明土壤呼吸对原始阔叶红松林人为干扰的响应是不同的。各林型Q10值介于1.84(人工落叶松林)—2.32(次生白桦林)之间。在整个生长季,各林型之间土壤呼吸的变异系数变化幅度为19.74%—37.39%,而各林型内土壤环间其变化幅度为32.13%—60.20%,显著大于样地间的变化幅度14.28%—35.70%(P<0.001),说明土壤呼吸在细微尺度上的差异更大。土壤湿度可以解释各林型(阔叶红松林除外)内部土壤呼吸15.8%—33.5%的空间异质性。     

长白山三种林型对蛾类群落结构和多样性的影响

选择长白山阔叶红松林、白桦林和落叶松林3种林型,对各林型蛾类的群落结构和多样性进行了比较。结果表明:蛾类各科(亚科)在3种林型中的比例分配明显不同。阔叶红松林中尺蛾科数量占优绝对优势,占47.0%,其次是舟蛾科,为9.3%。优势种为台褥尺蛾,占17.2%;从蛾类总种数和蛾类个体总数上看,白桦林中蛾类物种丰富度、多度和多样性指数明显高于阔叶红松林和落叶松林,优势度指数则最低;白桦林的优势种有尘尺蛾、枞灰尺蛾和双星白枝尺蛾;在落叶松林中,灯蛾科、波纹蛾科和枯叶蛾科的多度都比阔叶红松林和白桦林高,其优势种为阿泊波纹蛾。     

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.     

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.     

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。     

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.     

凉水自然保护区松鼠巢址选择的特征

2006 年6 月11 日至9 月27 日和2007 年8 月14 日至9 月21 日采用样线法,并结合定位观察的方法在凉水国家级自然保护区对松鼠的巢址选择进行了定量研究,共获取107 个巢样方和230 个对照样方。对每个样方分别测定了12 个巢树参数和7 个环境参数。统计分析结果表明,松鼠仅在针叶树上营巢,59.8 ± 4. 7% 的巢建于红松上,但对枝叶比红松更为浓密的云杉和臭冷杉有显著的选择性,而且显著倾向于在以云杉、臭冷杉为优势树种的生境中营巢。松鼠选择比较高大(树高21.5 ± 0.4 m)、活动通道比较多(3.6 ±0.1)的树营巢,巢树周围的树木密度(169 ± 4 株/ hm² )和多样性(辛普森指数0.51 ± 0.01)显著高于对照样方。巢址生境多位于南向的中、下坡位。巢址高12.4 ± 0.4 m,紧贴树干,多朝南。红松树上的巢多建于树冠的中下部,云杉和臭冷杉树上的巢多建于树冠的上部。松鼠巢对包括道路在内的林隙没有显著的回避。     

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 seed availability on habitat use by a specialist seed predator

Space-use patterns of seed predators are strongly affected by spatiotemporal variation in the abundance of different tree seeds, their major food source. However, most studies have measured relationships between overall food availability and space use, and there are few cases where effects of different food resources have been explored. We studied the effects of two food resources, Norway spruce and silver fir seeds, on space and habitat use in red squirrel in a subalpine conifer forest from 2000 to 2006. Fir seeds disperse in the autumn of the year they are produced, spruce the following spring. We estimated spruce and fir seed availability within individual home ranges and monitored home-range size using radiotelemetry. Males had larger home ranges than females and the sexes responded differently to variation in food and density. Spruce seed availability negatively affected home-range and core-area sizes of males in spring–summer. Space use was not affected by fir seed availability. Squirrels positively selected spruce for foraging and spruce was always preferred over fir. Our results showed that spruce, but not fir, affected space and habitat use of squirrels, suggesting they do not behaviourally respond to early seed dispersal in fir.     

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。