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凉水自然保护区贮食动物对红松种子的捕食与贮藏
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摘要
在2003年、2005年~2007年的秋季,我们在凉水自然保护区,对动物贮食红松(Pinus koreansis)种子行为进行了系统研究,结果显示:
     在凉水自然保护区,共有15种动物捕食红松种子,其中鸟类3目6科10种,哺乳动物2目3科5种,其中松鼠(Sciurus vulgaris)、星鸦(Nucifraga caryocatactes)、普通鳾(Sitta europaea)、花鼠(Eutamias sibiricus)4种动物具有分散贮食红松种子的行为。地表放置和埋藏实验显示,在凉水自然保护区内,动物对地表红松种子的捕食作用十分强烈,而埋藏能有效降低动物对红松种子的捕食。贮食动物对红松种子的贮藏一方面能有效降低种子的死亡率,另一方面也使大量的红松种子进入地被物下种子库,为红松天然更新储备了种源。
     针对普通鳾的贮食行为研究显示,其贮食行为谱由搬运行为、确定贮食点行为和贮藏行为组成,每个贮食点内的种子数均为1粒,贮食点平均深度为(1.4±0.5)cm,贮食微生境主要包括活立木、枯立木、倒木和倾斜地面。普通碍的贮点重取率接近100%。虽然单个普通碍的贮食能力明显弱于松鼠和星鸦,但由于保护区内普通鳾数量很大,故作为一个整体来看,其贮食竞争力还是比较可观的。而且普通鳾所贮藏的红松种子来源大多为松鼠、星鸦处理过的红松球果和贮点,故普通鳾和前两者之间存在一定的种间竞争。
     在凉水自然保护区,结实平年时,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,但与红松结实大年的贮点大小与深度之间存在显著差异。原始红松林内的贮食密度和贮食量最大。保护区内贮食动物的贮食总量约为118,000,00粒,其中松鼠的贮食量约为8,600,000粒,星鸦的贮食量约为3,100,000粒,松鼠在部分生境内的贮食密度和贮食量远大于星鸦,不考虑红松幼苗的适宜建成生境对红松天然更新的影响,单从贮藏量的角度,松鼠对红松天然更新种子库的贡献远大于星鸦。
     松鼠和星鸦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(Actinidiakolomikta)优势灌丛内贮藏红松种子,而星鸦优先选择在刺五加(Acanthopanaxsenticosus)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。
     松鼠和星鸦会选择具有萌发活力的红松种子贮藏在地被物下,在红松球果内种子只有73.2%可食用的情况下,松鼠和星鸦选择贮藏的红松种子100%饱满可食用,其贮藏行为有利于红松种子逃避其他动物的捕食。萌发实验表明,松鼠和星鸦贮点内的红松种子超过60%可萌发出红松幼苗,远远高于自然状态下红松种子的萌发率(41.7%)。在整个冬季贮食重取期内,松鼠会重取85%的贮点内的红松种子,星鸦会重取79%的贮点内的红松种子。对比圈养松鼠重取机制的结果,我们认为,在野外松鼠和星鸦主要依赖空间记忆能力和嗅觉重取其贮藏的红松种子,其中空间记忆能力起主要作用。
     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%。松鼠会将超过半数的红松球果搬运到300m以远的各种生境中进行贮藏,其贮食行为将有利于红松种群扩大分布区。
Synzoochory is a course of plants regeneration lead by hoarding behavior of animals. The research results indicate that frugivorous rodents are the main mammals providing synzoochory.More than 70 rodent species are known to have dispersal behavior benefit for the spread of plant seeds.The plant seeds dispersed by rodents include pignut,pine seeds, acorn etc.The animal species including frugivorous rodents formed stable diffuse coevolution relationship with plant species.The major researched of frugivorous rodent synzoochory developed three directions:describing and analyzing the synzoochory phenomena;exploring the hoarding behavioral mechanism;tracing the seeds fate after hoarding which is the most interested problem of researchers.The diplochory participated by multi-dispersal pattern in tropic and semi-tropic area is another studying highlight.The research efforts in future should be focused on improving the tracking technique of seeds fate,synzoochory research on ecosystem and landscape scale,discussing the effects of synzoochory in the ecosystem function and ecological restoration.
     The behavior of animals hoarding Korean Pine(Pinus koreansis) seeds was researched in autumn of 2003,2005~2007 in Liangshui Nature Reserves.
     There are 15 animals predating on Korean Pine seeds in Liangshui Nature Reserves,10 birds species of 6 families,3 orders,5 mammal species of 3 families 2 orders.Among them red squirrels(Sciurus vulgaris),Eurasian nutcracker(Nucifraga caryocatactes),Eurasian Nuthatch (Sitta europaea),chipmunk(Eutamias sibiricus) had behaviors of scatter-hoarding Korean Pine seeds.The ground placement and burying experiment revealed that the animals predated ground seeds intensively,and burying decreased the predation of animals on Korean Pine seeds effectively.
     Field research works of caching behavior by Eurasian Nuthatch in Liang-Shui Nature Reserve,Xiaoxing'anling Mt.Northeast China in October 16th to 27th.Caching behavior of Eurasian Nuthatch including carrying behavior,determining behavior and hoarding behavior. Average Caching time is(15.3±3.1) s,cache size is 1,cache depth is(1.4±0.5) cm. Caching microhabitat mainly including:live-stand wood,dead-stand wood,log and slope.
     The hoarding characteristic and cache quantity of Squirrels(Sciurus vulgaris) & Eurasian Nutcrackers(Nucifraga caryocatactes) were investigated in Liangshui Nature Reserve during September 30th to October 8th in 2005.The average cache size of squirrels was 3.02±0.1 and the depth was 2.67±0.08 cm;the average cache size and depth of Eurasian nutcrackers was 3.21±0.22 and 2.73±0.15 cm respectively.No significant difference in cache size and depth was showed between squirrels and Eurasian nutcrackers.The cache density and cache quantity by squirrels are most in primal Korean pine forest.The hoarding quantity of squirrels and Eurasian nutcrackers was 8 686 312±683 360 and 3 190 444±410 377 in the whole Nature Reserve yearly.The total hoarding quantity was 11 876 757±812 304.The hoarding behavior of squirrels and Eurasian nutcrackers is one of the main factors on natural regeneration of Korean pines.
     The habitat selection and microhabitat utilization for hoarding behavior of squirrels (Sciurus vulgaris) and Eurasian nutcrackers(Nucifraga caryocatactes) are investigated in Liangshui National Nature Reserve.Total 365 big samples and 1463 small samples were investigated in 11 forestry types.Results showed it is different selectivity of squirrels' hoarding behavior and Eurasian nutcrackers' hoarding behavior.The squirrels selected habitat order by major vegetation types is as follows:first natural spruce forest;second original Korean pine forest;third manmade Korean pine forest;fourth mix-conifer-broadleaf forest;fifth manmade spruce forest;sixth birch forest;seventh mix-conifer leaf forest;eighth manmade fallen leaves pine forest;ninth mix-broadleaf forest;tenth natural fir forest;finally other forest.The Eurasian nutcrackers selected habitat order by major vegetation types is as follows:first manmade Korean pine forest;second original Korean pine forest;third natural spruce forest;fourth manmade spruce forest;fifth mix-conifer-broadleaf forest;sixth mix-broadleaf forest;seventh birch forest;eighth mix-conifer leaf forest;ninth manmade fallen leaves pine forest;tenth natural fir forest;finally other forest. The Vanderploeg and Scavia selectivity index,Wi and Ei,were used to evaluate the microhabitat utilization selection of squirrels and Eurasian nutcrackers.Results indicated that their selections are sameness except brush type.
     Squirrels and Eurasian nutcrackers selected Korean Pine seeds with burgeoning activities. When there were73.2%available seeds in one cone the seeds hoarded by squirrels and Eurasian nutcrackers were esculent 100%.The burgeoning experiment revealed that 60%seeds in caches hoarded by squirrels and Eurasian nutcrackers can bud to seedlings with higher budding rate than that of nature status(41.7%).In winter squirrels recoveryed 85%caches of total caches,and Eurasian nutcrackers recoveryed 79%of total caches.
     In field the spatial memory ability and pilferer is the main tool of squirrels and Eurasian nutcrackers to recover cache.In little scale,squirrels mainly depend on pilferer to recover cache.In large scale,squirrels mainly depend on spatial memory ability to recover cache.All the factors to be concerned,spatial memory ability play a more important role than pilferer in cache recovery of squirrels in nature.
     The hoarding distance of Korean Pine(Pinus koreansis) seeds removed by squirrels (Sciurus vulgaris) was analyzed.The cones were marked by label,and observation in field was integrated to artificial placement experiments.The results indicated that Korean pine cones in different places were found and predated by squirrels at different time.The cones in disturbed places(near tourist path) were removed later.All cones disappeared within 10h. New cones(in 2006) and old cones(in 2005) were all predated and hoarded.Squirrels prefer to remove new cones and all the cones were hoarded,but old cones were eaten partly and only part cones were hoarded.Totally 110 marked cones were placed,in which 8 were eaten,102 were removed and hoarded,47 cones(46.1%) were found within 300m radial distance from placing spot.
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