农田林网中大斑啄木鸟对光肩星天牛控制作用的研究
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摘要
以光肩星天牛为主的杨树天牛是我国北方地区杨树林中最重要的害虫之一。上个世纪九十年代以来,杨树天牛的大规模发生给天敌鸟类提供了丰富的食物资源,大斑啄木鸟的种群数量有了明显的增长,并在一定程度上抑制了杨树天牛的发生和危害。本文以内蒙古乌拉特前旗地区的农田防护林作为研究对象,对该地区大斑啄木鸟的基础生物学和营养生态学进行了调查和分析,研究了大斑啄木鸟对光肩星天牛的控制作用,同时对农田林网条件下大斑啄木鸟的巢位和栖息地进行了深入的分析,并在此基础上开展了人工招引大斑啄木鸟技术研究。主要研究结果如下:
1.对农田林网条件下大斑啄木鸟的繁殖生物学和日间行为模式进行了深入的分析和研究,结果发现:大斑啄木鸟的繁殖期为4月上旬至6月中旬,日产卵一枚,窝卵数4-7枚,抱窝时长13d左右,孵化成功率约为51.52%,雏鸟第25天第一次飞离巢洞,育雏成功率为88.42%左右;觅食和啄食是大斑啄木鸟的主要活动行为,夏季具午间休息习性,日间行为节律明显,而冬季没有午休习性,日间行为节律不明显,不同性别大斑啄木鸟的日间行为差异不显著,但大斑啄木鸟在不同季节之间的日间行为差异极为显著,夏季活动时间明显长于冬季。
2.首次对农田林网条件下大斑啄木鸟的营养生态学进行系统的研究,明确了大斑啄木鸟不同季节的取食量、大斑啄木鸟对光肩星天牛的啄食率和啄食成功率,以及对取食对象和取食部位的选择性等,并对利用大斑啄木鸟控制光肩星天牛提出了合理的人为干扰措施。大斑啄木鸟春季、夏季和冬季的日取食次数分别为49.75±8.73次/天,57.75±8.68次/天和37.62±6.07次/天,对光肩星天牛幼虫的取食次数分别为1.58±0.27头/天,1.26±0.19头/天和11.91±1.92头/天,冬季是大斑啄木鸟控制光肩星天牛的主要季节;不考虑季节等其它因素的影响,大斑啄木鸟对光肩星天牛的啄食率为30.72%±29.06%,啄食成功率为88.33%±25.00%,不同季节的啄食率和啄食成功率存在一定的差异,且在枝干上的啄食次数明显高于主干,对小幼虫的啄食成功率明显高于大幼虫,但在觅食过程中,对幼虫大小没有明显的选择性。另外,大斑啄木鸟在枯死部位上的取食时间明显多于活立部位。根据以上调查结果,可以人为减少林地内的枯死木数量和清除枯死部位,增加大斑啄木鸟在活立部位的取食时间,从而提高对光肩星天牛的控制效果。
3.首次拟合出大斑啄木鸟对光肩星天牛的控制作用功能反应模型,明确了大斑啄木鸟对光肩星天牛的控制域值和控制效果。联合虫口密度和有虫株率可以很好地拟合出大斑啄木鸟对光肩星天牛的捕食作用功能反应模型,但不同季节的反应模型有所不同,冬季(夏季～冬季)和春季(冬季～春季)期间的捕食率与虫口密度和有虫株率呈反比关系,而夏季(春季～夏季)呈正比关系;以虫口密度计算,与对照样地相比,大斑啄木鸟对光肩星天牛的控制效果可以达到42.05%;对于不同光肩星天牛危害程度的样地,大斑啄木鸟的控制范围也不相同,轻度危害的林地1对大斑啄木鸟平均可以控制86.32±58.34公顷林地,中度危害的控制范围为53.51±12.99公顷,重度危害的控制范围为30.52±6.84公顷;在光肩星天牛的暴发期清除高虫口密度的被害木,可以有力地提高大斑啄木鸟对光肩星天牛的控制效果,实现对光肩星天牛的长期可持续控制目标。
4.首次对农田林网条件下大斑啄木鸟的巢位和栖息地进行系统的分析和研究,结果发现:大斑啄木鸟一年内有两个筑巢高峰,即入冬之前和春季育雏期,一年内平均每对大斑啄木鸟的新筑巢数为5.40±1.97个,影响其巢位选择的主要因子是树龄、树高、心腐程度、隐蔽度和避雨情况等,鸟巢的洞口方向主要分布于北向、东北和东向;影响大斑啄木鸟栖息地选择的主要因子是枯立木比例、有无片林、林地内主要树木的树龄、树高、胸径和食物丰富度等,平均每对大斑啄木鸟的栖息地面积为72.53±38.01公顷,变化范围为27.02～150.45公顷。
5.首次在农田林网条件下开展人工招引大斑啄木鸟研究,提出了人工招引大斑啄木鸟的技术要点。人工挂巢招引大斑啄木鸟应该在有大斑啄木鸟活动痕迹、枯立木少且林带整齐的林地内进行;挂巢时间在3月上旬之前,每年挂巢一次,每对大斑啄木鸟挂6～8个木段;巢木直径≥20cm,具有心腐和突起;挂巢高度3.5m左右,巢位间距可以设为100m左右,挂巢方向为北向、东北或者东向;木段应挂置在林带的中央,可以采用一字形或平行状排列。
Poplar longhorned beetles(PLB),with particular respect to Asian longhorned beetle Anoplophora glabripennis,are one of the most important groups of pest insects in the northern provinces of China.In these provinces the increase in PLB population density during the past three decades provided abundance of food supply for insectivorous birds,like woodpeckers,and caused the population increase of Great Spotted Woodpecker Picoides major since the beginning of 1990's.In plast years,reduction of PLB population was observed as a consequence of the higher feeding activity of P.major.In the present PhD dissertation,the biology and feeding ecology of P.major were investigated,as well as the control effect of P.major against A.glabripennis population.Moreover,an attraction technique using artificial nest-woods was established based on our experimental investigations of nest and habitat analysis of P. major in the agroforestry ecosystem of Wulateqianqi,Inner Mongolia from July 2004 to June of 2008. The main results achieved during this period were the following:
1.In the selected areas the breeding time of P.major was found to start from the beginning of April and to end in the middle of June.Females usually produced 4-7 eggs per one clutch,laying one egg per day.The duration of the incubation period was estimated around 13 days with about 51.2%eggs being hatched successfully.The nestling period was about 25 days before the young woodpecker flied away from the nest-cavity for the first time and the rate of breeding success was around 88.24%.
2.Foraging and pecking were the main behaviours of P.major observed during our investigations. Behavioural rhythm was clearly present in summer with a rest peak at midday,but a rhythm in behaviour was not recorded in winter.The time spent by the birds for movement was much longer in summer than in winter and significant differences were observed of behavioural patterns in different seasons.However,there was no detectable difference in behaviour between the two sexes.
3.Over the course of this study,foraging observations were recorded.The foraging times per day were 49.75±8.73,57.75±8.68,and 37.62±6.07 in spring,summer and winter respectively.The times per day when P.major pecked up A.glabripennis larvae during the three reported seasons were 1.58±0.27, 1.26±0.19 and 11.91±1.92,respectively.We found a significantly higher foraging on A.glabripennis larvae during the winter season compared to the other seasons(ANOVA:F=16.4010,df=2,P=0.0001). Therefore,winter was considered to be the crucial time to control A.glabripennis with P.major.The total annual rate of pecked A.glabripennis larvae by P.major was 30.72±29.06 and the rate of successful pecking was 88.33±25.00,not considering the impacting factors like different seasons and the age of A.glabripennis larvae,etc.However,the rate of pecked A.glabripennis larvae and rate of successful pecking were found to vary greatly during the different seasons,being 42.76±35.98,17.15±25.92 and 32.87±18.35 for rate of pecked larvae,and 89.76±18.43,74.00±42.53 and 94.94±11.10 for rate of successful pecking in spring,summer and winter respectively.The rate of successful pecking on young A.glabripennis larvae was much higher than that on old ones,but no detectable difference was observed between the rate of pecked larvae in different ages,that means no selection in the age of A. glabripennis larvae for P.major.The pecking times on branches was found to be more than on trunks, therefore we concluded that P.major prefers to forage from crown parts than from trunk parts of tree.In addition,P.major spent more time foraging on dead parts than on living parts of trees.According to these results,and to improve the effect of PLB population control,we decided to move away the dead trees and dead branches in order to push P.major foraging A.glabripennis larvae from the living parts of poplar trees.
4.The foraging function of P.major on the larval population of A.glabripennis was modelled, considering the population density of A.glabripennis and the rate of damaged trees.Different models were developed for each season.In winter and spring,the foraging rate of P.major was negatively related with the population density of A.glabripennis and with the rate of damaged trees,but positively related in summer.Autumn was not taken into consideration,due to the lack of sufficient investigations during this period.Comparing the investigated areas with control areas,where P.major was not present, we found that A.glabripennis population increased relatively slower of 42.05%due to the predation of P.major.The territory occupied by one breeding pair of P.major varies in function of the different population density of A.glabripennis,being 86.32±58.34 ha in slightly infested areas,53.51±12.99 ha in moderate infested areas and 30.52±6.84 ha in heavily infested areas..Measurements like moving away the severely damaged trees was estimated to be able to improve the control effect against A. glabripennis with P.major in long term.
5.Two nest-excavation peaks for P.major per year were observed.The first one occurred during late autumn-early winter and the second one occurred during the breeding time in spring.The average number of newly produced nests for one pair of P.major was 5.40±1.97 per year.The main affecting factors on the selection of nest sites were the age,height and rotten degree of tree and the concealment and protuberance for rain prevention of selected site.The orientation of nest-cavity entrance was found to be mainly directed to the north,north-east and east.The main affecting factors on the selection of habitats were the rate of dead trees,size of patch forest,age,height and diameter of tree and availability of food resource.The average area of habitat for one pair of P.major was estimated 72.53±38.01 ha, varying from 27.02 to 150.45 ha.
Based on these findings we propose that attraction with artificial nest-wood on P.major should be carried out in agroforests,where there were full shelter belts and P.major living.However,there should not be a high proportion of dead trees because P.major might prefer the natural nest-woods than the artificial ones.Artificial nest-woods should be hanged each year and the optimal hanging time was from end of February to beginning of March.The proper number of nest-wood was estimated to be 8 for one pair of P.major.Nest-woods should be rotten inside with the presence of clear protuberances and with a diameter larger than 20 cm.In our experiences,it is better to hang the nest-woods as high as possible, but practically the optimal height is around 3.5 m being really labour-costing to hang the nest-woods higher than 4.0 m.Nest-woods should be arranged in one line along the shelter belts with distance of 100 m between each other or in two paralleled lines.Nest-woods should be hanged on the host trees with the direction oriented to north,north-east or east.Moreover,the nest-woods should be hanged in the central part of an agroforest area to avoid disturbance and destruction.
1.对农田林网条件下大斑啄木鸟的繁殖生物学和日间行为模式进行了深入的分析和研究,结果发现:大斑啄木鸟的繁殖期为4月上旬至6月中旬,日产卵一枚,窝卵数4-7枚,抱窝时长13d左右,孵化成功率约为51.52%,雏鸟第25天第一次飞离巢洞,育雏成功率为88.42%左右;觅食和啄食是大斑啄木鸟的主要活动行为,夏季具午间休息习性,日间行为节律明显,而冬季没有午休习性,日间行为节律不明显,不同性别大斑啄木鸟的日间行为差异不显著,但大斑啄木鸟在不同季节之间的日间行为差异极为显著,夏季活动时间明显长于冬季。
2.首次对农田林网条件下大斑啄木鸟的营养生态学进行系统的研究,明确了大斑啄木鸟不同季节的取食量、大斑啄木鸟对光肩星天牛的啄食率和啄食成功率,以及对取食对象和取食部位的选择性等,并对利用大斑啄木鸟控制光肩星天牛提出了合理的人为干扰措施。大斑啄木鸟春季、夏季和冬季的日取食次数分别为49.75±8.73次/天,57.75±8.68次/天和37.62±6.07次/天,对光肩星天牛幼虫的取食次数分别为1.58±0.27头/天,1.26±0.19头/天和11.91±1.92头/天,冬季是大斑啄木鸟控制光肩星天牛的主要季节;不考虑季节等其它因素的影响,大斑啄木鸟对光肩星天牛的啄食率为30.72%±29.06%,啄食成功率为88.33%±25.00%,不同季节的啄食率和啄食成功率存在一定的差异,且在枝干上的啄食次数明显高于主干,对小幼虫的啄食成功率明显高于大幼虫,但在觅食过程中,对幼虫大小没有明显的选择性。另外,大斑啄木鸟在枯死部位上的取食时间明显多于活立部位。根据以上调查结果,可以人为减少林地内的枯死木数量和清除枯死部位,增加大斑啄木鸟在活立部位的取食时间,从而提高对光肩星天牛的控制效果。
3.首次拟合出大斑啄木鸟对光肩星天牛的控制作用功能反应模型,明确了大斑啄木鸟对光肩星天牛的控制域值和控制效果。联合虫口密度和有虫株率可以很好地拟合出大斑啄木鸟对光肩星天牛的捕食作用功能反应模型,但不同季节的反应模型有所不同,冬季(夏季～冬季)和春季(冬季～春季)期间的捕食率与虫口密度和有虫株率呈反比关系,而夏季(春季～夏季)呈正比关系;以虫口密度计算,与对照样地相比,大斑啄木鸟对光肩星天牛的控制效果可以达到42.05%;对于不同光肩星天牛危害程度的样地,大斑啄木鸟的控制范围也不相同,轻度危害的林地1对大斑啄木鸟平均可以控制86.32±58.34公顷林地,中度危害的控制范围为53.51±12.99公顷,重度危害的控制范围为30.52±6.84公顷;在光肩星天牛的暴发期清除高虫口密度的被害木,可以有力地提高大斑啄木鸟对光肩星天牛的控制效果,实现对光肩星天牛的长期可持续控制目标。
4.首次对农田林网条件下大斑啄木鸟的巢位和栖息地进行系统的分析和研究,结果发现:大斑啄木鸟一年内有两个筑巢高峰,即入冬之前和春季育雏期,一年内平均每对大斑啄木鸟的新筑巢数为5.40±1.97个,影响其巢位选择的主要因子是树龄、树高、心腐程度、隐蔽度和避雨情况等,鸟巢的洞口方向主要分布于北向、东北和东向;影响大斑啄木鸟栖息地选择的主要因子是枯立木比例、有无片林、林地内主要树木的树龄、树高、胸径和食物丰富度等,平均每对大斑啄木鸟的栖息地面积为72.53±38.01公顷,变化范围为27.02～150.45公顷。
5.首次在农田林网条件下开展人工招引大斑啄木鸟研究,提出了人工招引大斑啄木鸟的技术要点。人工挂巢招引大斑啄木鸟应该在有大斑啄木鸟活动痕迹、枯立木少且林带整齐的林地内进行;挂巢时间在3月上旬之前,每年挂巢一次,每对大斑啄木鸟挂6～8个木段;巢木直径≥20cm,具有心腐和突起;挂巢高度3.5m左右,巢位间距可以设为100m左右,挂巢方向为北向、东北或者东向;木段应挂置在林带的中央,可以采用一字形或平行状排列。
Poplar longhorned beetles(PLB),with particular respect to Asian longhorned beetle Anoplophora glabripennis,are one of the most important groups of pest insects in the northern provinces of China.In these provinces the increase in PLB population density during the past three decades provided abundance of food supply for insectivorous birds,like woodpeckers,and caused the population increase of Great Spotted Woodpecker Picoides major since the beginning of 1990's.In plast years,reduction of PLB population was observed as a consequence of the higher feeding activity of P.major.In the present PhD dissertation,the biology and feeding ecology of P.major were investigated,as well as the control effect of P.major against A.glabripennis population.Moreover,an attraction technique using artificial nest-woods was established based on our experimental investigations of nest and habitat analysis of P. major in the agroforestry ecosystem of Wulateqianqi,Inner Mongolia from July 2004 to June of 2008. The main results achieved during this period were the following:
1.In the selected areas the breeding time of P.major was found to start from the beginning of April and to end in the middle of June.Females usually produced 4-7 eggs per one clutch,laying one egg per day.The duration of the incubation period was estimated around 13 days with about 51.2%eggs being hatched successfully.The nestling period was about 25 days before the young woodpecker flied away from the nest-cavity for the first time and the rate of breeding success was around 88.24%.
2.Foraging and pecking were the main behaviours of P.major observed during our investigations. Behavioural rhythm was clearly present in summer with a rest peak at midday,but a rhythm in behaviour was not recorded in winter.The time spent by the birds for movement was much longer in summer than in winter and significant differences were observed of behavioural patterns in different seasons.However,there was no detectable difference in behaviour between the two sexes.
3.Over the course of this study,foraging observations were recorded.The foraging times per day were 49.75±8.73,57.75±8.68,and 37.62±6.07 in spring,summer and winter respectively.The times per day when P.major pecked up A.glabripennis larvae during the three reported seasons were 1.58±0.27, 1.26±0.19 and 11.91±1.92,respectively.We found a significantly higher foraging on A.glabripennis larvae during the winter season compared to the other seasons(ANOVA:F=16.4010,df=2,P=0.0001). Therefore,winter was considered to be the crucial time to control A.glabripennis with P.major.The total annual rate of pecked A.glabripennis larvae by P.major was 30.72±29.06 and the rate of successful pecking was 88.33±25.00,not considering the impacting factors like different seasons and the age of A.glabripennis larvae,etc.However,the rate of pecked A.glabripennis larvae and rate of successful pecking were found to vary greatly during the different seasons,being 42.76±35.98,17.15±25.92 and 32.87±18.35 for rate of pecked larvae,and 89.76±18.43,74.00±42.53 and 94.94±11.10 for rate of successful pecking in spring,summer and winter respectively.The rate of successful pecking on young A.glabripennis larvae was much higher than that on old ones,but no detectable difference was observed between the rate of pecked larvae in different ages,that means no selection in the age of A. glabripennis larvae for P.major.The pecking times on branches was found to be more than on trunks, therefore we concluded that P.major prefers to forage from crown parts than from trunk parts of tree.In addition,P.major spent more time foraging on dead parts than on living parts of trees.According to these results,and to improve the effect of PLB population control,we decided to move away the dead trees and dead branches in order to push P.major foraging A.glabripennis larvae from the living parts of poplar trees.
4.The foraging function of P.major on the larval population of A.glabripennis was modelled, considering the population density of A.glabripennis and the rate of damaged trees.Different models were developed for each season.In winter and spring,the foraging rate of P.major was negatively related with the population density of A.glabripennis and with the rate of damaged trees,but positively related in summer.Autumn was not taken into consideration,due to the lack of sufficient investigations during this period.Comparing the investigated areas with control areas,where P.major was not present, we found that A.glabripennis population increased relatively slower of 42.05%due to the predation of P.major.The territory occupied by one breeding pair of P.major varies in function of the different population density of A.glabripennis,being 86.32±58.34 ha in slightly infested areas,53.51±12.99 ha in moderate infested areas and 30.52±6.84 ha in heavily infested areas..Measurements like moving away the severely damaged trees was estimated to be able to improve the control effect against A. glabripennis with P.major in long term.
5.Two nest-excavation peaks for P.major per year were observed.The first one occurred during late autumn-early winter and the second one occurred during the breeding time in spring.The average number of newly produced nests for one pair of P.major was 5.40±1.97 per year.The main affecting factors on the selection of nest sites were the age,height and rotten degree of tree and the concealment and protuberance for rain prevention of selected site.The orientation of nest-cavity entrance was found to be mainly directed to the north,north-east and east.The main affecting factors on the selection of habitats were the rate of dead trees,size of patch forest,age,height and diameter of tree and availability of food resource.The average area of habitat for one pair of P.major was estimated 72.53±38.01 ha, varying from 27.02 to 150.45 ha.
Based on these findings we propose that attraction with artificial nest-wood on P.major should be carried out in agroforests,where there were full shelter belts and P.major living.However,there should not be a high proportion of dead trees because P.major might prefer the natural nest-woods than the artificial ones.Artificial nest-woods should be hanged each year and the optimal hanging time was from end of February to beginning of March.The proper number of nest-wood was estimated to be 8 for one pair of P.major.Nest-woods should be rotten inside with the presence of clear protuberances and with a diameter larger than 20 cm.In our experiences,it is better to hang the nest-woods as high as possible, but practically the optimal height is around 3.5 m being really labour-costing to hang the nest-woods higher than 4.0 m.Nest-woods should be arranged in one line along the shelter belts with distance of 100 m between each other or in two paralleled lines.Nest-woods should be hanged on the host trees with the direction oriented to north,north-east or east.Moreover,the nest-woods should be hanged in the central part of an agroforest area to avoid disturbance and destruction.
引文
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