白水江自然保护区蝶类多样性及珍稀种类濒危机制与保护措施研究
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摘要
本文在甘肃白水江自然保护区碧峰沟通过野外调查、定点定时观察、实验地饲养、标记重捕等方法研究了碧峰沟的蝶类多样性并进行了局部濒危评估；研究了四种重要蝶类的生物学特性；对长尾麝凤蝶的生境、自然种群生命表、种群结构、种群移动性进行了深入研究，并估测了各种种群参数，分析了种群发展趋势。分析了导致金裳凤蝶、长尾麝凤蝶种群下降或波动的关键因子，提出了保护措施。
1．碧峰沟蝶类多样性研究：研究了α-多样性和β-多样性。碧峰沟的物种多样性十分丰富。经调查，在30km~2的面积内，有蝶类180种，分属于12个科，97个属。β-多样性也很丰富，从沟口到沟底，植被破坏和污染逐渐减轻，蝶类种类多度和种群丰富度依次增加，β-多样性变化明显。
2．濒危评估：经评估，沟内蝶类濒危种类多。极危的种类有：美凤蝶、褐钩凤蝶、黑紫蛱蝶、大紫蛱蝶、斑凤蝶；濒危种类有：金裳凤蝶、箭环蝶、红基美凤蝶、牛郎凤蝶、突缘麝凤蝶、小黑斑凤蝶、波绿凤蝶、铂铠蛱蝶、傲白蛱蝶、猫蛱蝶；易危种类有：多姿麝凤蝶、乌克兰剑凤蝶、华夏剑凤蝶、宽带青凤蝶、冰清绢蝶、灰翅串珠环蝶。
3．重要种类的生物学、濒危原因与保护研究：
①金裳凤蝶：一年一代，以蛹在枝条上越冬。雄成虫5月下旬出现，雌成虫6月初出现，6月为盛期，7月下旬成虫少见。六月中旬产卵，卵期20天左右。幼虫期从六月下旬到九月中旬，幼虫五龄，历期50-55天，取食异叶马兜铃，八月中旬开始老熟幼虫在灌木、树枝上化蛹，蛹的颜色随化蛹时背景颜色变化，九月上旬为化蛹盛期。预蛹期2天，蛹期6个月。雄蝶飞翔能力强，飞翔高而快。雌蝶羽化后就近访花、交尾，寻找寄主植物马兜铃产卵、雌虫孕卵量为18-20粒；飞行缓慢，反应迟钝，相对容易捕捉。成虫活动时间在上午九点到下午六点，以上午10-12点最为活跃。
访花在羽化初期以合欢花为主，六月中旬后以粉叶羊蹄甲花为主，七月上旬后则以臭牡丹为主。雌雄性比为：1:3。
影响种群增长的主要原因是生境丧失、异常气候条件和天敌。生境的破坏、丧失减少了寄主植物的数量，夏季高温干旱、秋季淫雨降低了卵和幼虫的成活率，寄生蜂降低了越冬蛹的成活率。
卵和幼虫期的天敌主要有：蜘蛛、蠼螋、胡蜂、猎蝽、鸟类，蛹期天敌为姬蜂、鸟类。
种群结构为典型的异质种群，且为源-汇种群，共有九个斑块，山王庙、西沟斑块为最大的源种群斑块。种群在斑块内处于灭绝、再定居的动态过程，灭绝率大于再定植率，即种群处于下降趋势。
②小黑斑凤蝶：一年一代，以蛹越冬。翌年3月下旬至4月中旬羽化，幼虫绿色，
5龄。幼虫取食楠木叶，大龄幼虫常成排爬在向阳面叶片的正面晒太阳。因而容易被鸟
类捕食。蛹褐色，直形。
③嘉翠峡蝶:一年一代，以大龄幼虫或老熟幼虫在棕搁树棕中越冬。翌年五月中下
旬老幼虫从棕搁树棕中爬出，取食嫩棕叶。6月上旬老熟幼虫化蛹，蛹期20天。7月一8
月为成虫期。9月至10月为幼虫期，10月底以幼虫在棕搁树棕中越冬。成虫喜在日光
下活动，飞翔迅速，行动活泼;卵黑褐色，有瘤状突起;幼虫绿色，体两侧长满长刺;
蛹为悬蛹，绿色，背部有5个亮丽的银色金属斑点。
卵和幼虫由于裸露在棕叶背面，易被天敌吃掉，存活率低。卵孵化率为50%，幼虫
存活率为49.1%。
④长尾庸凤蝶:一年二代，以蛹在灌木或树枝上越冬，翌年4月中旬羽化。第一
代卵历期12一14天，第二代卵历期7一9天。第一代幼虫5龄，历期30天;第二代幼虫5
龄或6龄，历期30一40天;幼虫取食异叶马兜铃。第一代蛹6月初化蛹，蛹期20一26天;
第二代蛹7月下旬到9月底化蛹。第一代成虫5月中下旬为高峰期;第二代成虫高峰期
在6月下旬到7月中旬。有世代重叠。雄成虫比雌成虫早羽化7一10天，主要在沟底活
动，飞行能力较强。雌成虫飞行能力较差，主要在出生地附近访花交尾产卵。产卵量平
均为31.5粒。雌雄性比1:4。雄成虫寿命26天，雌成虫寿命21天。成虫访花植物有:
广布野豌豆、兰花、火棘、合欢、粉叶羊蹄甲、臭牡丹、密蒙花、接骨草。
种群增长率参数为:第一代:越冬蛹羽化率:55.1%;卵孵化率:75石%;幼虫保
存率:42.1%;第二代:蛹羽化率:83.3%;卵孵化率:53.0%;幼虫保存率:22.8%。
种群增长率:第一代成虫到第二代成虫:2.021;第二代成虫到越冬后第一代成虫:0.508。
年增长率为:1.027。
种群数量:用标记重捕法调查了4代成虫的种群数量，结果为:2001年第二代:n83
头;2002年第一代:586头;2002年第二代:1211头;2003年第一代:6巧头;
环境容量为:第一代:600头:第二代:1 200头。
种群密度:第一代:20头/枷2;第二代:40头加mZ。
最小居住时间:雄成虫最长14天，平均4.6天;雌成虫最长20天，平均6.6天。
家域的确定:用至少捕到两次的数据确定。
种群结构:为典型的异质种群，且为源一汇种群，一共有九个斑块，山王庙、西沟斑
块为最大的源种群斑块。种群在斑块内没有灭绝发生，即种群处于在环境容量附近上下
波动的平衡状态。
_种群移动性:雄成虫扩散距离在Zkm以内的个体比例89.19%;扩散距离在3kl
This study deals with butterfly diversity and assessing endangered species in Natural Reserve of Baishuijiang in Gansu Province. The research includes field investigation, observation at certain time and places, captive breeding in experimental area, capture-recapture etc. The biology of four important butterfly species is studied, especially focusing on the natural population life table, habitats, population structure and mobility of Byasa impediens. The population developing trend and key factor that result population decline and fluctuation of Troides aeacus and Byasa impediens are analysed. Conservation measure are provided.
1. Butterfly diversity in Bifeng Gully
This work includes research on a and P -diversity. It appears that the species hi this area are very abundant. There are 180 butterfly species in 30km2, which belong to 12 families and 97 genera. Meanwhile, ?-diversity is also in high level with gradually increasing abundance and diversity when pollution and damage of forestry are decreasing from the mouth to the end of this gully.
2. The assessing of endangered degree of some species
There are many endangered species in this gully and the critically endangered species include Papilio epycides, Meandrusa scrion, Sasakia funebris.Sasakia charonda, Chilasa clytia ; the endangered species include Papilio alcmenor, Papilio bootes, Papilio polyctor, Byasa plutonius, Chilasa epycides, Chitoria pallas, Jimelaea maculata, Helcyra superba,Stichophthalma howqua, and the vulnerable species include Byasa polyeutes, Pazala tamerlanta, Pazala mandarina, Pamassius latreille, Graphium scopoli, Funis aepope.
3. The biology, endangered reason and conservation measure of important species
(1) Troides aeacus: univoltine, overwintering as pupae on branch. Male adults fly in late May and females emerge in early June. The adults are seldom found in late July. Female lays its eggs hi mid-June and eggs will hatch in 20 days, larvae feeding on Aristolochia kaempferi from late June to mid-September (about 50-55 days), five instars. From mid-August, larvae become pupae in tree branches or brushes with changeable color based on background. The pre-pupa is about 2 days and pupa is about six months. The male adults can fly higher and faster than females. Adults will visit flowers, mating and find hostplant Aristolochia kaempferi after eclosion. The active time of adults is from 9:00 A.M. to 18:00 P.M., more active from 10-12 A.M. in the morning. The rate of female to male is 1:3.
Their favorable flower is Albizzia julibrissin in the early eclosing period, and then change to Aristolochia kaempferi in mid-June, then Clerodendrum bungei after early July.
The key factors, which influence the growth of population of Troides aeacus, include habitat loss, abnormal climatic conditions and natural enemies. The loss of habitat results in less hostplants. The hot and dry weather in summer and too much rain in autumn reduce the survival rate of eggs and larvae rapidly. The natural enemies of larvae include spiders, earwigs, wasps, bugs and birds, and pupa ichneumon.
The population structure is typical metapopulation (source-sink population) with nine patches. The patch of West Gully and Shanwang-Temple Gully is the source population and the population in these patches is a dynamic process from extinction to reclonizition. The rate of extinction is higher than reclonizition, which indicates the decline of population.
(2) Chilasa epycides: Univoltine. They overwinter in pupae and eclose in next late March to Mid-April. Larvae are greenish, five instars, feeding on leaves of Aristolochia kaempferi. The larvae of late instars always array in lines on the leaves exposed to the sun to catch the heat of light, so they are easily to be found and eaten by birds. The pupa of this is brown.
(3) Euthalia kardama: Univoltine. They overwinter in elder larvae in the old leaves of palms in late October. The larvae emerge from the leaves in mid-to late- May of next year and feed on fresh leaves of palms. The larvae will become
1．碧峰沟蝶类多样性研究：研究了α-多样性和β-多样性。碧峰沟的物种多样性十分丰富。经调查，在30km~2的面积内，有蝶类180种，分属于12个科，97个属。β-多样性也很丰富，从沟口到沟底，植被破坏和污染逐渐减轻，蝶类种类多度和种群丰富度依次增加，β-多样性变化明显。
2．濒危评估：经评估，沟内蝶类濒危种类多。极危的种类有：美凤蝶、褐钩凤蝶、黑紫蛱蝶、大紫蛱蝶、斑凤蝶；濒危种类有：金裳凤蝶、箭环蝶、红基美凤蝶、牛郎凤蝶、突缘麝凤蝶、小黑斑凤蝶、波绿凤蝶、铂铠蛱蝶、傲白蛱蝶、猫蛱蝶；易危种类有：多姿麝凤蝶、乌克兰剑凤蝶、华夏剑凤蝶、宽带青凤蝶、冰清绢蝶、灰翅串珠环蝶。
3．重要种类的生物学、濒危原因与保护研究：
①金裳凤蝶：一年一代，以蛹在枝条上越冬。雄成虫5月下旬出现，雌成虫6月初出现，6月为盛期，7月下旬成虫少见。六月中旬产卵，卵期20天左右。幼虫期从六月下旬到九月中旬，幼虫五龄，历期50-55天，取食异叶马兜铃，八月中旬开始老熟幼虫在灌木、树枝上化蛹，蛹的颜色随化蛹时背景颜色变化，九月上旬为化蛹盛期。预蛹期2天，蛹期6个月。雄蝶飞翔能力强，飞翔高而快。雌蝶羽化后就近访花、交尾，寻找寄主植物马兜铃产卵、雌虫孕卵量为18-20粒；飞行缓慢，反应迟钝，相对容易捕捉。成虫活动时间在上午九点到下午六点，以上午10-12点最为活跃。
访花在羽化初期以合欢花为主，六月中旬后以粉叶羊蹄甲花为主，七月上旬后则以臭牡丹为主。雌雄性比为：1:3。
影响种群增长的主要原因是生境丧失、异常气候条件和天敌。生境的破坏、丧失减少了寄主植物的数量，夏季高温干旱、秋季淫雨降低了卵和幼虫的成活率，寄生蜂降低了越冬蛹的成活率。
卵和幼虫期的天敌主要有：蜘蛛、蠼螋、胡蜂、猎蝽、鸟类，蛹期天敌为姬蜂、鸟类。
种群结构为典型的异质种群，且为源-汇种群，共有九个斑块，山王庙、西沟斑块为最大的源种群斑块。种群在斑块内处于灭绝、再定居的动态过程，灭绝率大于再定植率，即种群处于下降趋势。
②小黑斑凤蝶：一年一代，以蛹越冬。翌年3月下旬至4月中旬羽化，幼虫绿色，
5龄。幼虫取食楠木叶，大龄幼虫常成排爬在向阳面叶片的正面晒太阳。因而容易被鸟
类捕食。蛹褐色，直形。
③嘉翠峡蝶:一年一代，以大龄幼虫或老熟幼虫在棕搁树棕中越冬。翌年五月中下
旬老幼虫从棕搁树棕中爬出，取食嫩棕叶。6月上旬老熟幼虫化蛹，蛹期20天。7月一8
月为成虫期。9月至10月为幼虫期，10月底以幼虫在棕搁树棕中越冬。成虫喜在日光
下活动，飞翔迅速，行动活泼;卵黑褐色，有瘤状突起;幼虫绿色，体两侧长满长刺;
蛹为悬蛹，绿色，背部有5个亮丽的银色金属斑点。
卵和幼虫由于裸露在棕叶背面，易被天敌吃掉，存活率低。卵孵化率为50%，幼虫
存活率为49.1%。
④长尾庸凤蝶:一年二代，以蛹在灌木或树枝上越冬，翌年4月中旬羽化。第一
代卵历期12一14天，第二代卵历期7一9天。第一代幼虫5龄，历期30天;第二代幼虫5
龄或6龄，历期30一40天;幼虫取食异叶马兜铃。第一代蛹6月初化蛹，蛹期20一26天;
第二代蛹7月下旬到9月底化蛹。第一代成虫5月中下旬为高峰期;第二代成虫高峰期
在6月下旬到7月中旬。有世代重叠。雄成虫比雌成虫早羽化7一10天，主要在沟底活
动，飞行能力较强。雌成虫飞行能力较差，主要在出生地附近访花交尾产卵。产卵量平
均为31.5粒。雌雄性比1:4。雄成虫寿命26天，雌成虫寿命21天。成虫访花植物有:
广布野豌豆、兰花、火棘、合欢、粉叶羊蹄甲、臭牡丹、密蒙花、接骨草。
种群增长率参数为:第一代:越冬蛹羽化率:55.1%;卵孵化率:75石%;幼虫保
存率:42.1%;第二代:蛹羽化率:83.3%;卵孵化率:53.0%;幼虫保存率:22.8%。
种群增长率:第一代成虫到第二代成虫:2.021;第二代成虫到越冬后第一代成虫:0.508。
年增长率为:1.027。
种群数量:用标记重捕法调查了4代成虫的种群数量，结果为:2001年第二代:n83
头;2002年第一代:586头;2002年第二代:1211头;2003年第一代:6巧头;
环境容量为:第一代:600头:第二代:1 200头。
种群密度:第一代:20头/枷2;第二代:40头加mZ。
最小居住时间:雄成虫最长14天，平均4.6天;雌成虫最长20天，平均6.6天。
家域的确定:用至少捕到两次的数据确定。
种群结构:为典型的异质种群，且为源一汇种群，一共有九个斑块，山王庙、西沟斑
块为最大的源种群斑块。种群在斑块内没有灭绝发生，即种群处于在环境容量附近上下
波动的平衡状态。
_种群移动性:雄成虫扩散距离在Zkm以内的个体比例89.19%;扩散距离在3kl
This study deals with butterfly diversity and assessing endangered species in Natural Reserve of Baishuijiang in Gansu Province. The research includes field investigation, observation at certain time and places, captive breeding in experimental area, capture-recapture etc. The biology of four important butterfly species is studied, especially focusing on the natural population life table, habitats, population structure and mobility of Byasa impediens. The population developing trend and key factor that result population decline and fluctuation of Troides aeacus and Byasa impediens are analysed. Conservation measure are provided.
1. Butterfly diversity in Bifeng Gully
This work includes research on a and P -diversity. It appears that the species hi this area are very abundant. There are 180 butterfly species in 30km2, which belong to 12 families and 97 genera. Meanwhile, ?-diversity is also in high level with gradually increasing abundance and diversity when pollution and damage of forestry are decreasing from the mouth to the end of this gully.
2. The assessing of endangered degree of some species
There are many endangered species in this gully and the critically endangered species include Papilio epycides, Meandrusa scrion, Sasakia funebris.Sasakia charonda, Chilasa clytia ; the endangered species include Papilio alcmenor, Papilio bootes, Papilio polyctor, Byasa plutonius, Chilasa epycides, Chitoria pallas, Jimelaea maculata, Helcyra superba,Stichophthalma howqua, and the vulnerable species include Byasa polyeutes, Pazala tamerlanta, Pazala mandarina, Pamassius latreille, Graphium scopoli, Funis aepope.
3. The biology, endangered reason and conservation measure of important species
(1) Troides aeacus: univoltine, overwintering as pupae on branch. Male adults fly in late May and females emerge in early June. The adults are seldom found in late July. Female lays its eggs hi mid-June and eggs will hatch in 20 days, larvae feeding on Aristolochia kaempferi from late June to mid-September (about 50-55 days), five instars. From mid-August, larvae become pupae in tree branches or brushes with changeable color based on background. The pre-pupa is about 2 days and pupa is about six months. The male adults can fly higher and faster than females. Adults will visit flowers, mating and find hostplant Aristolochia kaempferi after eclosion. The active time of adults is from 9:00 A.M. to 18:00 P.M., more active from 10-12 A.M. in the morning. The rate of female to male is 1:3.
Their favorable flower is Albizzia julibrissin in the early eclosing period, and then change to Aristolochia kaempferi in mid-June, then Clerodendrum bungei after early July.
The key factors, which influence the growth of population of Troides aeacus, include habitat loss, abnormal climatic conditions and natural enemies. The loss of habitat results in less hostplants. The hot and dry weather in summer and too much rain in autumn reduce the survival rate of eggs and larvae rapidly. The natural enemies of larvae include spiders, earwigs, wasps, bugs and birds, and pupa ichneumon.
The population structure is typical metapopulation (source-sink population) with nine patches. The patch of West Gully and Shanwang-Temple Gully is the source population and the population in these patches is a dynamic process from extinction to reclonizition. The rate of extinction is higher than reclonizition, which indicates the decline of population.
(2) Chilasa epycides: Univoltine. They overwinter in pupae and eclose in next late March to Mid-April. Larvae are greenish, five instars, feeding on leaves of Aristolochia kaempferi. The larvae of late instars always array in lines on the leaves exposed to the sun to catch the heat of light, so they are easily to be found and eaten by birds. The pupa of this is brown.
(3) Euthalia kardama: Univoltine. They overwinter in elder larvae in the old leaves of palms in late October. The larvae emerge from the leaves in mid-to late- May of next year and feed on fresh leaves of palms. The larvae will become
引文
1．中国科学院生物多样性委员会，《生物多样性研究的原理与方法》，1994，北京：中国科学出版社237
2．Richard B．Primack著，祁承经译，1996《保护生物学概论》，长沙：湖南科学技术出版社，351页
3．宋延龄等，《物种多样性研究与保护》，1998，浙江科学技术出版社261页．
4．Richard Primack，季维智，《保护生物学基础》，2000，北京，中国林业出版社
5．徐汝梅，《昆虫种群生态学》，1987，北京：北京师范大学出版社409页
6．吴千红，邵则信，《昆虫生态学实验》，1991，上海，复旦大学出版社
7．袁志发，周静芋，《试验设计与分析》，2000，北京，高等教育出版社
8．周尧，《中国蝶类志》1999，洛阳：河南科学技术出版
9．白九维，王效岳，中国凤蝶志，1998，淑馨出版社
10．赵力，王效岳，中国鳞翅目(3)，1997，台湾省立博物馆
11．刘灿然，马克平，生物多样性的测度方法ⅵ：与多样性测度有关的统计问题，生物多样性1998，6(3)：229-239。
12．陈振宁，曾阳，青海祁连地区不同生境类型蝶类多样性研究，生物多样性2001，9(2)：109-114
13．赵士洞，任毅，“国际生物多样性观察年(IBOY)”核心项目——内容介绍及建议，2001，9(2)：196-200
14．许宏发，陆厚基，最小存活种群(MVP)——保护生物学的一个基本理论Ⅰ．最小存活种群及其在野生动物保护中的应用1996，15(2)：25-30
15．许宏发，陆厚基，最小存活种群——保护生物学的一个基本理论Ⅱ．物种灭绝的过程和最小存活种群(种群脆弱性分析PVA)生态学杂志 1996，15(3)：50-55
16．许宏发，陆厚基，王小明，玛他种群：种群生态学理论应用于保护生物学实践的新范例，生态学杂志，1998，17(1)：47-53
17．王文，分子系统学在生物保护中的意义，生物多样性，1998，6(2)：138-142
18．陈小勇，历宁，Meta-种群结构及其遗传后果，生态学杂志 2000，19(5)：71-75
19．胡萃，吴小晶，王选民，珍稀濒危昆虫——中华虎凤蝶的生物学，昆虫学报 1992，35(2)：195-199
20．袁德成等，1998 中华虎风蝶栖息地、生物学和保护现状，生物多样性，1998，6(2)：105-115
21．文礼章，陈永年，潘洞，豆芽实验种群生命表参数的研究，昆虫学报 1991，34(1)：26-31
22．吴子江，薛智华，张治，棉铃虫自然种群生命表的研究及其应用，生态学报 1993，13(2)：185-193
23．渠春梅，韩兴国，苏波，片断化森林的边缘效应与自然保护区的设计管理，生态学报，2000，20(1)
24．解焱，国际濒危物种等级新标准，生物多样性，1995，3(4)：234-239
25. Amy Knight, Lincoln P. Brower Fls, Ernest H. Williams, Spring Remigration of The Monarch
26． Butterfly, Danaus plexippus (Lepidoptera: Nymphglidae) in North-Central Florida: Estimating Population Parameters Using Mark-recapture,Biology Journal of the Society, 1999,68:531-556． With 7 figures
27． Chtfles J. Krebs 1998 f33Ecological Methodology f44 Second Edition Addison-Welsey Educational Publishers,Inc. 1-580．
28． C. Stelter, J. Settele and C. Wissel ,1996, Die Bedeutung von Stoerungen und Pflegemassnahmen fuer das Ueberdauern von Schmetterlingspopulationen im Kontext eines Modells. Verh. Ges. Oekologie 26, 483-483
29． Chris D. Thomas, Catastrophic Extinction of Population Sources in a Butterfly Meta-Population, The American Naturalist 1996, Vol. 148,No.6．
30． C. D. Thomas, Spatial and Temporal Variability in a Butterfly Population, Oecologia ,1991, 87:577-580．
31． C. D. Thomas, 1992 Distribution of Occupied and Vacant Butterfly Habitats in Fragmented Landscapes, Oecologia 92:563-567．
32． C. D. Thomas & J.C.G. Abery, Estimating Rates of Butterfly Decline from Distribution Maps :the effect of Scale, Biological Conservation ,1995,73:59-65．
33． C. D. Thomas, J. A.Thomas, and M. S. Warren, Distributions of Occupied and Vacant Butterfly Habitats in Fragmented Landscapes, Oecilogia ,1992,92:563-567
34． C. D. Thomas, Extinction, Colonization, and Metapopulations: Enviromental Tracking by Rare Species, Conservation Biology, 1994,8(2) :373-378
35． Cheryl B. Schultz, Dispersal Behavior and its Implications Oregon Butterfly , Conservation Biology, 1998,Vol.i2,No.2:284-292
36． Cheiyl B. Schultz , Dispersal Behavior and Its Implications for Reserve Design in Rare Oregon Butterfly, Conservation Biology, 1998,12(2) :284-292
37． Dennis D. Murphy, An Environment Metapopulation Approach to Population Viability analysis for a Threatened Invertebrate, Conservation Biology, 1990,Vol.4,No.l
38． Denis F. Owen, Larval Food Shortage and Adult Dispersal in Callimorpha dominula (L.) (Lepidoptera: Arctiidae), Nota lepid. 1996,18(3/4) :281-288
39． E. Pollard and T. J. Yates 1993 (Monitoring Butterflies for Ecology and Conservation) .
40． Graeme Caughley Anne Gunn 1996 (Conservation Biology in Theory and Practice) Blackwell Since.Inc. 1-458．
41． H. Resit Akcakaya,RAMAS/metapop,Viability Analysis for Stage-structured Metapopulations,User Manual for version 1． 0
42． Hugh B. Britten, Pefer F. Brussard,Dennis D. Murphy, The Pending Extinction of Uncompahgre Fritlary Butterfly, Conservation Biology, 1994,8(1) :86-94
43． Hanski ,Gilpin 1997 Academic Press 1-454．
44． Ilkka Hanski, Spatial Scale, Patchiness and Population Dynamics on Land, Phil. Trans. R. Soc. Land B (1994) 343,19-25
45． Ilkka Hanski, Mikko Kuussaari, and Marko Nieminen, Metepopulation Structure and Migration in The Butterfly Melitaea Cinxia,Ecology, 1994, 75(3) :747-762
46． Illka Hanski and Mats Gyllenberg, Uniting Two General Patterns in the Distribution of Species, Science,1997,vol.275
47． Illka Hanski, Chris D. Thomas, Metapopulation dynamics and Conservation: a Spatially Explicit Model Applied to Butterflies, Biological Conservation 68 (1994) 167-180
48． Illka Hanski, Atte Moilanen, and Mats Gyllenberg, Minimum Viable Metapopulation Size, The American Naturalist ,1996, vol. 147,No.4
49． Illka Hanski, Juha Poeyry, Timo Pakkala & Mikko Kuussaari, Multiple Equilibria in metapopulation Dynamics,Nature, 1995,337(19) :618-621
50． Illka Hanski, Timo Pakkala, Mikko Kuussaari and Guangchun Lei, Metapopulation Persistence of an Endangered Butterfly in a Fragmented landscape, oijos 72:21-28． Copenhagen,1995
51． Illka Hanski, A Practical Model of Metapopulation Dynamics, Journal of Animal Wcology 1994,63,151-162
52． Illka Hanski, The Quantitative Incidence Function Model and Persistence of an Endangered Butterfly Metapopulation, Conservation Biology , 1996,10(2) :578-590
53 . Illka Hanski, Patch-Occupancy Dynamics in Fragmented Landscapes, tree , 1994,9(4) : 131-135
54． Josef Settele 1999 (Die Tagfalter Deutschlands > erlag Eugen Ulmer GmbH.
55． Josef Settele, Reinart Feldmann.Klaus Henle, Populationsgroessenschaetzung bei Tieren, Ausgewaehlte Verfahren fuer den Einsatz in Populationsoekologie und Naturschutz-Kurzfassung einer Internet-Version, Naturschutz und Landschaftsplanung,Zeitschrift fuer angewandte Oekologie,Heft 6/1998,Juni 30． Jahegang.
56． Josef Settele, Reinart FeIdmann,Klaus Henle, Populationsgroessenscrfaetzung bei Tieren, Ausgewaehlte Verfahren fuer den Einsatz in Populationsoekologie und Naturschutz-Kurzfassung einer Internet-Version, Naturschutz und Landschaftsplanung,Zeitschrift fuer angewandte Oekologie,Heft 6/1998,Juni 30． Jahegang.
57． Josef Settele 1996 < Species Survival in Fragmented Landscapes> Klwwer Academic Publishhers 1-381．
58． J. Settele, R. Pauler and K. Kockelke, 1995 Magerrasennutzung und Anpassungen bei Tagfaltern : Populationsoekologische Forschung als Basis fuer Schutzmassnahmen am Beispiel von Glaucopsyche (Maculinea) arion (Thymian-Ameisenblaeuling) und Glaucopsyche (Mmaculined) rebeli (Kreuzenzian-Ameisenblaeuling). In.B.Beinlich and H.Plachter (Hrsg.):Schutz und Entwicklung der
Kalkmagerrasen der Schwaebischen Alb. Beihefte zu den Veroeffentlichungen fuer Naturschutz und Landschaftspflege in Baden-Wuerttemberg 83, 129-158
59． J. Settele, and H. J. Poethke 19% Towards demographic population vulnerability categories of butterflies. In: J. Settele, C. Margules, P. Poschlod and K. Henle (eds.). Species Survival in Fragmented Landscapes. Kluwer, Dordrecht, 282-289
60． J. Settele, R. feldmann, K. Henle, K. Kockelke and H.-J. Poethke 1998 Populations-groessenschaetzung bei Tieren-ausgewaehlte Verfahren fuer den Einsatz in Populationsoekologie und Naturschutz (Kurzfassung einer Internet-Version), landschaftsplanung 30(6) , 174-181
61． J. Settele, R. Feldmann , K. Henle , K. Kockelke and H.-J. Poethke 1998 Populations-groessenschaetzung bei Tieren-ausgewaehlte Verfahren fuer den Einsatz in Populationsoekologie und Naturschutz (Internet-Version). Internet-Ablage zum vorgenannten Manuskript (mit auefuehrlicher Textversion und Programmen) unter: 'http://www.ufz.de/spb/nat/settele.html'(36pp)
62． J. Settele, K. Henle and C. Bender ,1996 Metapopulationen und Biotopverbund: Theorie und Praxis am Beispiel von Schmetterlingen und Reptilien. Zeitschrift flier Oekologie und Naturschutz 5, 187-206
63． J. Settele, Reinart Feldmann, Klaus Henle, Methoden der quantitativen Erfassung von Tagfaltern, 1999 《Die Tagfalter Deutschlands 》 erlagEugenUlmer GmbH. 145
64． J. A. Thomas ,1991 Rare Species Conservation :Case Studies of European Butterflies.
65． J. A. Thomas and M. G Morris 1994 , Patterns Mechanisms and Rates of Extinction Among Invertebrates in the united kingdom, Phil. Trans. R.Soc. Lond. B: 344:47-54．
66． J. A. Thomas , The ecology and conservation of Maculinea arion and other European Species of large blue butterfly , Ecology and Conservation of Butterflies, 1995,Chapman &Hall.
67． J. A. Thomas, A Quick Method for Estimating Butterfly Numbers During Surveys, Biological Conservation, 1983 ,27: 195-211 .
68． J. A. Thomas, M. L. Munguira , J. Martin and G W. Elmes , Basal Hatching by Maculinea butterfly eggs: a Consequence of Advanced Myrmecophily? Biological Journal of the linnean society, 1991,44:175-184,with 3 figures.
69． J. A. Thomas, J. C. Wardlaw, The Effect of Queen Ants on The Survival of Maculinea arion Larvae in Myrmica ant nests, Oecologia,1990,85:87-91
70． Karin Amler Josef Settele 1999 Verlag Eugen Ulmer GmbH.
71． Klaus Fisher, Population Structure, Mobility and Habitat Selection of the Butterfly Lycaena Hippothoe ( Lycaenidae: Lycaenini) in Western Germany, nota lepid.1998 ,21(1) :14-30;
72． Katrin Vogel 1997, Sonne Ziest und Flockenblmen Was braucht eine ueberlebensfaehrige
Population des Roten Seheckenfalters (Melitaea clidyma) , Dissertation zur Erlangung des Naturwissenschaflichen Doltorgrades der Universitaet Wuerzburg.
73． Leonard Nunney, 1994 Estimating the Effective Population Size of Conserved Populations , Conservation Biology, Vol.8,No. 1:175-184
74． MichaelJ. Samways 1995 Chapman & Hall 1-357．
75． Mark L. Sharffer, Population Viability Analysis, Conservation Biology,1990,4(l):39-40
76． Manfred Alban pfeifer, Ulf Robert Andrick, Wolfgang Frey & Jossef Settele, on The ethology and Ecology of a Small and Isolated population of The Dusky Large Blue Butterfly Glaucopsyche (Maculined) nausithous (Lycaenidae), Nota lepid. 2000, 23(2) : 147-172;
77． N.Mark Collins 1985 IUCN Gland Switzerland and Cambridge U.K.
78． Pauler-Fuerste, R. GK aule and J. Settele 1996 Aspects of the population vulnerability of the large blue butterfly, Glaucopsyche (Maculinea) arion, in south-west Germany. In: J. Settele, C. Margules, P. Poschlod and K. Henle (eds.). Species Survival in Fragmented Landscaes. Kluwer, Dordrecht, 275-281．
79． Paul R. Ehrlich, Dennis D. Murphy, Conservationg Lessens from Long-Term Studies of Checkerspot Butterflies, Conservation Biology , 1987,1(2) :122-131
80． Richard B. Harris, Lynn A. Maguire, Mark L. Shaffer, Sample Sizes for Minimum Viable Population Estimation, Conservation Biology, 1987, 1 ( 1 ):72-75
81． Ron Panzer and Mark W. Schwartz, Effective of a Vegetation-Based Approach to Insect Conservation, 1998 , Conservation Biology, Vol.l2,No.3:693-702
82． R Feldmann, K. Henle and J. Settele 1995 Applicability of mark-recapture methods to butterfly conservation research. In: Bissonette, J. A. and P. R. Krausman (eds.): Integrating people and wildlife for a sustainable future. Proceedings of the first International Wildlife Management Congress. The Wildlife Society, Bethesda,Md.,624-626
83． Roland Steiner , Habitatnutung, Arealdynamik und Schutzaspekte einer Population des Segelfalters, Diplomarbeit der Fakaltaet fuer Biologie Universitaet Hohenheim.
84． Soule 1987 (Viable Populations for Conservation) Cambridge University Press.
85． Soule 1986 Conservation Biology SinauerAssociates.Inc.
86． T. R. E. South wood & P. A. Henderson Ecological Methods Third Edition.
87． scale, Biological Conservation 73:59-65．
88． T. R. E. Southwood & P. A. Henderson Ecological Methods .Third Edition.
89． Yan-ling Song, Population Viability Analysis for Two Isolated Populations of Hainan Elds Deer, Conservation Biology, Pages 1467-1472,Volume 10,No.5,October 1996．
2．Richard B．Primack著，祁承经译，1996《保护生物学概论》，长沙：湖南科学技术出版社，351页
3．宋延龄等，《物种多样性研究与保护》，1998，浙江科学技术出版社261页．
4．Richard Primack，季维智，《保护生物学基础》，2000，北京，中国林业出版社
5．徐汝梅，《昆虫种群生态学》，1987，北京：北京师范大学出版社409页
6．吴千红，邵则信，《昆虫生态学实验》，1991，上海，复旦大学出版社
7．袁志发，周静芋，《试验设计与分析》，2000，北京，高等教育出版社
8．周尧，《中国蝶类志》1999，洛阳：河南科学技术出版
9．白九维，王效岳，中国凤蝶志，1998，淑馨出版社
10．赵力，王效岳，中国鳞翅目(3)，1997，台湾省立博物馆
11．刘灿然，马克平，生物多样性的测度方法ⅵ：与多样性测度有关的统计问题，生物多样性1998，6(3)：229-239。
12．陈振宁，曾阳，青海祁连地区不同生境类型蝶类多样性研究，生物多样性2001，9(2)：109-114
13．赵士洞，任毅，“国际生物多样性观察年(IBOY)”核心项目——内容介绍及建议，2001，9(2)：196-200
14．许宏发，陆厚基，最小存活种群(MVP)——保护生物学的一个基本理论Ⅰ．最小存活种群及其在野生动物保护中的应用1996，15(2)：25-30
15．许宏发，陆厚基，最小存活种群——保护生物学的一个基本理论Ⅱ．物种灭绝的过程和最小存活种群(种群脆弱性分析PVA)生态学杂志 1996，15(3)：50-55
16．许宏发，陆厚基，王小明，玛他种群：种群生态学理论应用于保护生物学实践的新范例，生态学杂志，1998，17(1)：47-53
17．王文，分子系统学在生物保护中的意义，生物多样性，1998，6(2)：138-142
18．陈小勇，历宁，Meta-种群结构及其遗传后果，生态学杂志 2000，19(5)：71-75
19．胡萃，吴小晶，王选民，珍稀濒危昆虫——中华虎凤蝶的生物学，昆虫学报 1992，35(2)：195-199
20．袁德成等，1998 中华虎风蝶栖息地、生物学和保护现状，生物多样性，1998，6(2)：105-115
21．文礼章，陈永年，潘洞，豆芽实验种群生命表参数的研究，昆虫学报 1991，34(1)：26-31
22．吴子江，薛智华，张治，棉铃虫自然种群生命表的研究及其应用，生态学报 1993，13(2)：185-193
23．渠春梅，韩兴国，苏波，片断化森林的边缘效应与自然保护区的设计管理，生态学报，2000，20(1)
24．解焱，国际濒危物种等级新标准，生物多样性，1995，3(4)：234-239
25. Amy Knight, Lincoln P. Brower Fls, Ernest H. Williams, Spring Remigration of The Monarch
26． Butterfly, Danaus plexippus (Lepidoptera: Nymphglidae) in North-Central Florida: Estimating Population Parameters Using Mark-recapture,Biology Journal of the Society, 1999,68:531-556． With 7 figures
27． Chtfles J. Krebs 1998 f33Ecological Methodology f44 Second Edition Addison-Welsey Educational Publishers,Inc. 1-580．
28． C. Stelter, J. Settele and C. Wissel ,1996, Die Bedeutung von Stoerungen und Pflegemassnahmen fuer das Ueberdauern von Schmetterlingspopulationen im Kontext eines Modells. Verh. Ges. Oekologie 26, 483-483
29． Chris D. Thomas, Catastrophic Extinction of Population Sources in a Butterfly Meta-Population, The American Naturalist 1996, Vol. 148,No.6．
30． C. D. Thomas, Spatial and Temporal Variability in a Butterfly Population, Oecologia ,1991, 87:577-580．
31． C. D. Thomas, 1992 Distribution of Occupied and Vacant Butterfly Habitats in Fragmented Landscapes, Oecologia 92:563-567．
32． C. D. Thomas & J.C.G. Abery, Estimating Rates of Butterfly Decline from Distribution Maps :the effect of Scale, Biological Conservation ,1995,73:59-65．
33． C. D. Thomas, J. A.Thomas, and M. S. Warren, Distributions of Occupied and Vacant Butterfly Habitats in Fragmented Landscapes, Oecilogia ,1992,92:563-567
34． C. D. Thomas, Extinction, Colonization, and Metapopulations: Enviromental Tracking by Rare Species, Conservation Biology, 1994,8(2) :373-378
35． Cheryl B. Schultz, Dispersal Behavior and its Implications Oregon Butterfly , Conservation Biology, 1998,Vol.i2,No.2:284-292
36． Cheiyl B. Schultz , Dispersal Behavior and Its Implications for Reserve Design in Rare Oregon Butterfly, Conservation Biology, 1998,12(2) :284-292
37． Dennis D. Murphy, An Environment Metapopulation Approach to Population Viability analysis for a Threatened Invertebrate, Conservation Biology, 1990,Vol.4,No.l
38． Denis F. Owen, Larval Food Shortage and Adult Dispersal in Callimorpha dominula (L.) (Lepidoptera: Arctiidae), Nota lepid. 1996,18(3/4) :281-288
39． E. Pollard and T. J. Yates 1993 (Monitoring Butterflies for Ecology and Conservation) .
40． Graeme Caughley Anne Gunn 1996 (Conservation Biology in Theory and Practice) Blackwell Since.Inc. 1-458．
41． H. Resit Akcakaya,RAMAS/metapop,Viability Analysis for Stage-structured Metapopulations,User Manual for version 1． 0
42． Hugh B. Britten, Pefer F. Brussard,Dennis D. Murphy, The Pending Extinction of Uncompahgre Fritlary Butterfly, Conservation Biology, 1994,8(1) :86-94
43． Hanski ,Gilpin 1997
44． Ilkka Hanski, Spatial Scale, Patchiness and Population Dynamics on Land, Phil. Trans. R. Soc. Land B (1994) 343,19-25
45． Ilkka Hanski, Mikko Kuussaari, and Marko Nieminen, Metepopulation Structure and Migration in The Butterfly Melitaea Cinxia,Ecology, 1994, 75(3) :747-762
46． Illka Hanski and Mats Gyllenberg, Uniting Two General Patterns in the Distribution of Species, Science,1997,vol.275
47． Illka Hanski, Chris D. Thomas, Metapopulation dynamics and Conservation: a Spatially Explicit Model Applied to Butterflies, Biological Conservation 68 (1994) 167-180
48． Illka Hanski, Atte Moilanen, and Mats Gyllenberg, Minimum Viable Metapopulation Size, The American Naturalist ,1996, vol. 147,No.4
49． Illka Hanski, Juha Poeyry, Timo Pakkala & Mikko Kuussaari, Multiple Equilibria in metapopulation Dynamics,Nature, 1995,337(19) :618-621
50． Illka Hanski, Timo Pakkala, Mikko Kuussaari and Guangchun Lei, Metapopulation Persistence of an Endangered Butterfly in a Fragmented landscape, oijos 72:21-28． Copenhagen,1995
51． Illka Hanski, A Practical Model of Metapopulation Dynamics, Journal of Animal Wcology 1994,63,151-162
52． Illka Hanski, The Quantitative Incidence Function Model and Persistence of an Endangered Butterfly Metapopulation, Conservation Biology , 1996,10(2) :578-590
53 . Illka Hanski, Patch-Occupancy Dynamics in Fragmented Landscapes, tree , 1994,9(4) : 131-135
54． Josef Settele 1999 (Die Tagfalter Deutschlands > erlag Eugen Ulmer GmbH.
55． Josef Settele, Reinart Feldmann.Klaus Henle, Populationsgroessenschaetzung bei Tieren, Ausgewaehlte Verfahren fuer den Einsatz in Populationsoekologie und Naturschutz-Kurzfassung einer Internet-Version, Naturschutz und Landschaftsplanung,Zeitschrift fuer angewandte Oekologie,Heft 6/1998,Juni 30． Jahegang.
56． Josef Settele, Reinart FeIdmann,Klaus Henle, Populationsgroessenscrfaetzung bei Tieren, Ausgewaehlte Verfahren fuer den Einsatz in Populationsoekologie und Naturschutz-Kurzfassung einer Internet-Version, Naturschutz und Landschaftsplanung,Zeitschrift fuer angewandte Oekologie,Heft 6/1998,Juni 30． Jahegang.
57． Josef Settele 1996 < Species Survival in Fragmented Landscapes> Klwwer Academic Publishhers 1-381．
58． J. Settele, R. Pauler and K. Kockelke, 1995 Magerrasennutzung und Anpassungen bei Tagfaltern : Populationsoekologische Forschung als Basis fuer Schutzmassnahmen am Beispiel von Glaucopsyche (Maculinea) arion (Thymian-Ameisenblaeuling) und Glaucopsyche (Mmaculined) rebeli (Kreuzenzian-Ameisenblaeuling). In.B.Beinlich and H.Plachter (Hrsg.):Schutz und Entwicklung der
Kalkmagerrasen der Schwaebischen Alb. Beihefte zu den Veroeffentlichungen fuer Naturschutz und Landschaftspflege in Baden-Wuerttemberg 83, 129-158
59． J. Settele, and H. J. Poethke 19% Towards demographic population vulnerability categories of butterflies. In: J. Settele, C. Margules, P. Poschlod and K. Henle (eds.). Species Survival in Fragmented Landscapes. Kluwer, Dordrecht, 282-289
60． J. Settele, R. feldmann, K. Henle, K. Kockelke and H.-J. Poethke 1998 Populations-groessenschaetzung bei Tieren-ausgewaehlte Verfahren fuer den Einsatz in Populationsoekologie und Naturschutz (Kurzfassung einer Internet-Version), landschaftsplanung 30(6) , 174-181
61． J. Settele, R. Feldmann , K. Henle , K. Kockelke and H.-J. Poethke 1998 Populations-groessenschaetzung bei Tieren-ausgewaehlte Verfahren fuer den Einsatz in Populationsoekologie und Naturschutz (Internet-Version). Internet-Ablage zum vorgenannten Manuskript (mit auefuehrlicher Textversion und Programmen) unter: 'http://www.ufz.de/spb/nat/settele.html'(36pp)
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