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1、松鼠贮食生境选择对红松天然更新的影响宗诚1 戎可1,2 马建章基金项目：国家自然科学基金面上项目 (30470235 ) ; 内蒙古自治区高等学校科学研究项目(NJzy08164)作者简介：宗诚(1973 -),吉林盘石人,博士,副教授,主要从事野生动物生态教学和研究工作.Email: * 通讯作者,corresponding author, E-mail: 致谢：野外工作得到凉水国家级自然保护区管理局全体员工的大力支持，英文摘要的修改得到比利时安特卫普大学Lucas A. Wauters教授的帮助，谨致谢意Foundation item: The project was financiall
2、y supported by National Natural Science Foundation of China (No. 30470235), University Research Project of Inner Mongolia (No. NJzy08164).Biography: ZONG Cheng, Ph. D, Associate professor, mainly engaged in animal ecology. E-mail: * 程志奔1(1 东北林业大学野生动物资源学院, 哈尔滨 150040) (2 呼伦贝尔学院生命科学与化学学院, 海拉尔 021008)摘
3、要：以松鼠贮食红松种子时丢弃的塔核为指征，在凉水国家级自然保护区16和19林班内对松鼠的贮食生境选择进行研究。在369hm2范围内，均匀布设10条总长15km的样线，收集有效样方343个。根据对样方内塔核数量的聚类分析结果判断，其中贮食样方184个，对照样方159个。对贮食样方11个生境因子的主成分分析结果显示，种源距离、林型、红松幼苗数量、灌丛密度和类型是影响松鼠贮食生境选择的主要因子。使用Baileys方法分析表明：1.松鼠对针叶混交林、云冷杉林、人工云杉林具有选择倾向性；2.对人工红松林、人工落叶松林则表现为回避；3.对天然红松林表现为随机利用。150-600m的种源距离对松鼠的贮食生境
4、选择没有影响。50%以上的松塔被丢弃在距离红松母树300m以内的红松林林缘和红松林林隙中，显著高于300m以外(One-Way ANOVA Test; df=3,183, F=5.76, p=0.0009)，表明松鼠可以将塔核携带出红松母树林。贮食样方内的灌木密度显著低于对照样方(Kruskal-Wallis test; df=1, 2=83.99, p0.0001)。贮食样方内红松幼苗数量显著高于对照样方(Kruskal-Wallis test; df=1, 2=104.13, p0.0001)，说明松鼠的贮食生境适于红松更新苗的建成。松鼠贮食红松种子对于红松种群的更新和扩散具有促进作用。关
5、键词：松鼠; 红松; 贮食行为; 生境选择; 塔核The effect of habitat selection of hoarding on natural regeneration of the Korean pines by the Eurasian red squirrels, Sciurus vulgarisZONG Cheng1, RONG Ke1,2, MA Jian-Zhang1* , CHENG Zhi-Ben11 College of Wildlife Resources, Northeast Forest University, Harbin 150040,China2
6、School of Life Science and chemistry, Hulunbeier College, Hailar 021008, ChinaAbstract: We used discarded cone-cores, left by Eurasian red squirrels, to study the habitat selection of Korean pine-seeds hoarding, in compartments No.16 and No.19 in Liangshui Nature Reserve, China. Ten lines, in total
7、of 15km long, were uniformly collocated, and 343 valid samples were collected in a 369hm2 area. One hundred and eighty-four were hoarding samples while the other 159 were control samples according to the cluster analysis based on the number of the cone-cores. The principal component analysis, using
8、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 Baileys method indicated that the squirrels showed 1.pr
9、eference for natural coniferous forest, natural fir and spruce forest and planted spruce forest; 2.avoidance of planted Korean pine forest and planted larch forest; 3.random use of natural Korean pine forest Moreover the distance from the Korean pines in the range of 150-600m showed no effect on the
10、 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 less than 300m than at 300m away (One-Way ANOVA Test; df=3,183, F=5.76, p=0.0009). This demonstrated that the E
11、urasian red squirrels could took 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, 2=83.99, p0.0001). The number of the Korean pine seedlings in samples of hoarding area was
12、 significantly higher than that in the control samples (Kruskal-Wallis test; df=1, 2=104.13, p0.0001). 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
13、the regeneration and dispersal of the Korean pines.Key Words: Eurasian red squirrels (Sciurus vulgaris); Korean pine (Pinus koraiensis); hoarding behavior; habitat selection; cone-core阔叶红松林是小兴安岭生态系统的顶级群落，对于维护我国东北的生态安全具有特殊的意义1。由于种子的结构特点，红松(Pinus koraiensis)的天然更新对动物有近乎绝对的依赖性，贮食动物为越冬贮藏的红松种子成为红松天然更新的种源2
14、, 3。松鼠(Sciurus vulgaris)作为分散贮食(scatter hoarding)动物，秋季大量贮藏红松种子作为越冬的主要食物4。然而，松鼠在协助红松天然更新过程中的地位却存在争议，Hutchins（1996）和鲁长虎（2001）认为松鼠不具有促进红松扩散的生态意义3, 5，马建章(2006)的研究则得出不完全相同的结论6。要确定松鼠是否能够促进红松扩散，要解决两个问题：1)松鼠能否将红松种子贮藏于母树林外；2)松鼠的贮食生境是否有利于红松更新苗的建成。马建章(2006)主要从贮点微生境特征的角度部分回答了这两个问题，本研究在从较大尺度上以松鼠贮食红松种子时丢弃的塔核和红松更新苗
15、为指征，分析松鼠的贮食生境选择特征，以判断松鼠能否促进红松种群的更新与扩散。1 研究地概况野外研究地为黑龙江省凉水国家级自然保护区，保护区概况参见此前报道4, 6。具体研究地点位于凉水自然保护区内相邻的16号和19号两个林班，总面积369hm2。在这两个林班内分布有凉水自然保护区所有典型的森林类型，而且受人类松塔采集活动的影响相对较小。2 研究方法2.1 调查方法根据2003年以来对野外松鼠贮食行为的观察，松鼠丢弃在贮点附近的塔核(cone-core)比较完整，与其他动物丢弃的塔核有明显区别，可以作为松鼠贮食地点的标志，丢弃的塔核数量也可以间接反映松鼠贮食量6。2008年10月15日11月10
16、日，在凉水自然保护区16和19林班沿东西方向均匀布设10条样线，每条样线长度1.5km，样线间距150m，覆盖16和19林班全境所有不同的林型。样线上每隔50m布设一个半径10m的圆形样方，每遇林型改变即增设一个样方，共调查样方344个。分别计数每个样方内松鼠丢弃塔核数量，测量林型、针叶树数量、灌丛类型、灌丛密度、地面覆盖物类型、郁闭度、坡向、坡位、红松幼苗数量、与红松母树林关系、种源距离等11个生境因子，具体方法如下：林型分为原始红松林、针叶混交林、阔叶混交林、针阔混交林、次生白桦林、云冷杉林、人工红松林、人工落叶松林和人工云杉林9种6。分别将上述林型依次以自然数1-9赋值；针叶树数量指样方内包括红松、云杉、冷杉在内的针叶树的株数；灌丛类型以样方内占优势的灌丛种类命名，分为无灌
