笼养珍稀鹳类（Ciconiidae）的行为生态学研究
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摘要
本文以国家Ⅰ级保护动物笼养白鹳和黑鹳为研究对象，采用瞬间和扫
描取样法（instantaneous and scan sampling）进行连续时间采样（time
sampling），结合全事件记录法（all occurrence sampling），记录其行为类型。
统计所得数据利用SPSS10.0 for Windows及Statistica软件包进行统计分
析。
建立笼养条件下白鹳（下称白鹳）和黑鹳目前较详尽的行为谱，为对该
物种的进一步的行为学研究奠定了基础。详细阐述了繁殖行为过程。记述
了人工条件下白鹳和黑鹳的幼体行为及身体发育的过程，并对其身体各组
成部分的变化拟合了生长发育模型。
研究结果表明在自身内源节律和外界环境及人为因素综合影响下，笼
养白鹳和黑鹳的各种行为在1天中和1年中表现出一定的节律性、周期性。
笼养条件下白鹳和黑鹳的各种行为节律受饲喂方式影响明显，固定的投食
时间使取食行为节律与野外个体不同，进而影响其他行为类型，加上其内
源节律及气候因子的影响，从而出现人工条件下特有的行为节律。在年行
为节律中，繁殖活动对其影响较大，气候因子特别是气温有较明显的影响。
由于笼舍大小的限制和种群规模小，白鹳的时间分配是以满足自身需
要为主，其社群行为较少。另外在笼养条件下，充足易得的食物供应及相
对稳定安全的人工环境，使白鹳和黑鹳减少了觅食和警戒的时间，从而表
山东师范大学硕士研兜失学位论文
现出取食时间较少而休息时间较长的时间分配模式；并且在繁殖期间，通
常会明显地改变时间分配。不同生理状态下的群体间时间分配出现了不同
的差异。
行为多样性指数可作为衡量同种或同域分布近缘种间对外界环境适
应性大小或存活能力的指标。结果表明白鹤的不同性别、繁殖状态和不同
年龄影响其行为多样性指数，并且白鹤的行为多样性指数显著高于黑鹤。
行为多样性指数在I年中具有显著的周期性，繁殖活动对其有显著影响。
可根据行为多样性指数针对各种情况对其进行不同的护理。
在人工环境下，对白鹤和黑鹤影响较大的环境因子主要有人类活动。
气温和湿度等。人工环境对于幼年个体较成年个体影响更大，因此，是否
创建适宜的人工环境关系到迁地保护的成败。
The behaviors of Oriental white storks Ciconia ctconia bOyciana and
black Storks Ciconia nlera were observed by instamaneous and scan samPling
and a1l occurrence sampling method, and the variety of behavior of Storks
were registered. Data were Statistical inference by the software of SPSS10.0
fOr Wndows and Statistica.
The most detailed and a11-round behavior ethogram of Orienial white
stork and B1ack stork by now was recorded, and made a prepare for further
ethologic reseaxch in other day The behavior of breeding procedUre was
expatiated at laxge. And the development of behavior and body of squabs of
Orienta white stork and Black Stork were recorded and the eqUaional models
were regressive analysed.
The resu1ts are as fol1ows: With the effect of endogenCtic rhythm and
environment and artificial factors, the behavor of Oriental White stork and
black stork in caPtivity has a dai1y and annual periodicity The behavior
thythms of storks were affected by feeding way remarkably. It was fixed time
of feeding, Which made the feeding behavior rhythm of storks in caPtivity
different from that of wt1d storks, that wtl1 triher effect the others behavior
rhy'thm, in addition to the effect of tdrie of feeding, endogenetic thythm and
environment factors also contribute to the behavior the in caPtivity
specifical1y Breeding behavior had some distinct dripact on annua1 rhythm of
behaviof, and weather factors, or rather air temPeratW also had an obvious1y
effect.
Because of Iimit of aviny and the litt1e population of beVy the activities
of se1fmaintenance was first in the time budgets of the Oriental White storks
4
in captive, the other maintenance was second. Sex, age, and breeding or not
had some impact on time budgets. The enough fOod provided and the
quiescent circumstance made the Storks short the time to feeding and more
time to have rest.
Behavior diversity index can be used as a spobol to value adaPtive ability.
The reseaxch results show that sex, age, and breeding or not had some impact
on. behavior diversity index. The behavior diversity index of Oriental white
stork is 1axger than that of Black stork. The index appeared periodicity in one
year's time, and activities of breeding have significan1y influence on the
periodicity
In anificia1 eniironmeni, the most importam environmental factors that
effect Oriental white Stork and Black stork are activities of human, air
temperatUre and hUmidity. The artificial entironment have a more sighfican
impact in particular to squabs than to adults.
描取样法（instantaneous and scan sampling）进行连续时间采样（time
sampling），结合全事件记录法（all occurrence sampling），记录其行为类型。
统计所得数据利用SPSS10.0 for Windows及Statistica软件包进行统计分
析。
建立笼养条件下白鹳（下称白鹳）和黑鹳目前较详尽的行为谱，为对该
物种的进一步的行为学研究奠定了基础。详细阐述了繁殖行为过程。记述
了人工条件下白鹳和黑鹳的幼体行为及身体发育的过程，并对其身体各组
成部分的变化拟合了生长发育模型。
研究结果表明在自身内源节律和外界环境及人为因素综合影响下，笼
养白鹳和黑鹳的各种行为在1天中和1年中表现出一定的节律性、周期性。
笼养条件下白鹳和黑鹳的各种行为节律受饲喂方式影响明显，固定的投食
时间使取食行为节律与野外个体不同，进而影响其他行为类型，加上其内
源节律及气候因子的影响，从而出现人工条件下特有的行为节律。在年行
为节律中，繁殖活动对其影响较大，气候因子特别是气温有较明显的影响。
由于笼舍大小的限制和种群规模小，白鹳的时间分配是以满足自身需
要为主，其社群行为较少。另外在笼养条件下，充足易得的食物供应及相
对稳定安全的人工环境，使白鹳和黑鹳减少了觅食和警戒的时间，从而表
山东师范大学硕士研兜失学位论文
现出取食时间较少而休息时间较长的时间分配模式；并且在繁殖期间，通
常会明显地改变时间分配。不同生理状态下的群体间时间分配出现了不同
的差异。
行为多样性指数可作为衡量同种或同域分布近缘种间对外界环境适
应性大小或存活能力的指标。结果表明白鹤的不同性别、繁殖状态和不同
年龄影响其行为多样性指数，并且白鹤的行为多样性指数显著高于黑鹤。
行为多样性指数在I年中具有显著的周期性，繁殖活动对其有显著影响。
可根据行为多样性指数针对各种情况对其进行不同的护理。
在人工环境下，对白鹤和黑鹤影响较大的环境因子主要有人类活动。
气温和湿度等。人工环境对于幼年个体较成年个体影响更大，因此，是否
创建适宜的人工环境关系到迁地保护的成败。
The behaviors of Oriental white storks Ciconia ctconia bOyciana and
black Storks Ciconia nlera were observed by instamaneous and scan samPling
and a1l occurrence sampling method, and the variety of behavior of Storks
were registered. Data were Statistical inference by the software of SPSS10.0
fOr Wndows and Statistica.
The most detailed and a11-round behavior ethogram of Orienial white
stork and B1ack stork by now was recorded, and made a prepare for further
ethologic reseaxch in other day The behavior of breeding procedUre was
expatiated at laxge. And the development of behavior and body of squabs of
Orienta white stork and Black Stork were recorded and the eqUaional models
were regressive analysed.
The resu1ts are as fol1ows: With the effect of endogenCtic rhythm and
environment and artificial factors, the behavor of Oriental White stork and
black stork in caPtivity has a dai1y and annual periodicity The behavior
thythms of storks were affected by feeding way remarkably. It was fixed time
of feeding, Which made the feeding behavior rhythm of storks in caPtivity
different from that of wt1d storks, that wtl1 triher effect the others behavior
rhy'thm, in addition to the effect of tdrie of feeding, endogenetic thythm and
environment factors also contribute to the behavior the in caPtivity
specifical1y Breeding behavior had some distinct dripact on annua1 rhythm of
behaviof, and weather factors, or rather air temPeratW also had an obvious1y
effect.
Because of Iimit of aviny and the litt1e population of beVy the activities
of se1fmaintenance was first in the time budgets of the Oriental White storks
4
in captive, the other maintenance was second. Sex, age, and breeding or not
had some impact on time budgets. The enough fOod provided and the
quiescent circumstance made the Storks short the time to feeding and more
time to have rest.
Behavior diversity index can be used as a spobol to value adaPtive ability.
The reseaxch results show that sex, age, and breeding or not had some impact
on. behavior diversity index. The behavior diversity index of Oriental white
stork is 1axger than that of Black stork. The index appeared periodicity in one
year's time, and activities of breeding have significan1y influence on the
periodicity
In anificia1 eniironmeni, the most importam environmental factors that
effect Oriental white Stork and Black stork are activities of human, air
temperatUre and hUmidity. The artificial entironment have a more sighfican
impact in particular to squabs than to adults.
引文
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88 Bryant D M,Tatner P, 1988.Energetics of the annual cycle of dippers Cinclus cinclus[J].Ibis, 130:17～38.
89 Morton M L,1967.Diural feeding patterns in white-crowned sparrows Zonotrichia leucophrys gambelii[J].The Conder, 69:491～512.
90 Verbeek N A M, 1972.Daily andannual time budget of the yellow-billed magpie[J].Auk, 89:576～582.
91 杨晓君，文贤继，王淑珍，杨岚等，2000．笼养大紫胸鹦鹉取食活动。动物学研究 21(2)：115～120。
92 杨晓君，文贤继，杨岚，1994．笼养白腹锦鸡觅食活动的观察[J]。动物学研究，15(1)：49～54。
93 杨晓君，文贤继，杨岚，1995。笼养白腹锦鸡繁殖期的时间分配[J]。动物学研究，16 (2)：178～184。
94 杨晓君，周建平，杨岚，1991．棕胸竹鸡觅食行为的观察[J]。动物学研究，12(3)：259～264。
95 杨晓君，杨岚，1996。笼养绿孔雀行为活动时间分配的初步观察[J]。动物学报，42(增刊)：106～111。
96 田蕴，高玮，相桂权等，1996．人工饲养条件下花尾榛鸡的行为[J]。动物学报，42(增刊)：31～38。
97 杨晓君，杨岚，王淑珍，文贤继，1998．笼养大紫胸鹦鹉的活动时间分配[J]。动物学报，44 (3)：277～285。
98 Quinlan,E.E.&Baldassare,G.A. 1984. Activity budgets of nonbreeding green-winged teal on playa lakes in Texas. J.Wildl. Mgmt 48:838～845.
99 Bryant, C.J.& Thompson,D.B.A.1985.Gulls and Plovers: the ecology and behavior of mixed-species feeding groups. Beckenham:Croom Helm.
100 Carmi-Winkler, N.,Degen,A.A.&Pinshow, B. 1987. Seasonal time-energy budgets of free-livingchukars in the Negev desert. Condor 89:594～601.
101 Caraco,T. 1979. Time budgeting and group size: aTest of theory. Ecology 60:618～627.
102 East,M. 1980. Sex differences and the effect of temperature on the foraging behaviour of Robins Erithacus rubecula. Ibis 122:517～520.
103 East, M. 1982. Time-budgeting by European robins Erithacus rubecula: inter and intrasexual comparisons during autumn,winter and early spring. Ornis Scand 13:85～93.
104 Pearson O P, 1954.The daily energy requirements of a wild Anna Hummingbird. The Conder,56:317～322.
105 Hixon,M.A.,Carpenter, F.L.&Paton,D.C. 1983. Territory area,flower density, and time budgeting in hummingbirds:an experimental and theorectical anaslysis. Am.Nat. 122:366～391.
106 Gibb,J. 1956. Food,feeding habits and territory of the Rock Pipit Anthus spinoletta. Ibis 98: 506～530.
107 Hunter, M.L.,jr.,Witham,J.W.&Dow, H. 1984. Effects of a carbyl-induced depression in invertebrate abundance on the growth and behavior of American black duck and mallard ducklings. Can. J. Zool. 62:452～456.
108 Davies,N.B.&Lundberg,A. 1984. Food distribution and a variable mating system in the dunnock ,Prunella modularis. J.A nim. Ecol.53:895～912.
109 Enoksson,B.1990 Autumn territories and population regulation in the nuthatch Sitta europaea: An experimental study. J.Anim.
110 Ydenberg,R.C. 1984. The conflict between feeding and territorial defence in the great tit. Behav. Ecol. Sociobiol.15:103～108.
111 Davies,N.B.&Lundberg,A. 1985. The influence of food on time budgets and timing of breeding in the Dunnock Prunella modularis. Ibis 127:100～110.
112 Verner, J.1965. Time budget of the male long-billed marsh wren during the breeding season. Condor 67:125～139.
113 Maxson,S.J.&Oring,L.W. 1980. Breeding season time and energy budgets of the polyandrous spotted sandpiper. Behaviour 74:200～263.
114 Turner, A.K. 1982. Timing of laying by swallows (Hirundo rustica) and sand martins (Riparia riparia). J.Anim. Ecol. 51:29～46.
115 Siegfried,W.R. 1974. Time budgets of behavior among lesser scaups on Delta Marsh. J. Wildl. Mgmt 38:708～713.
116 Widen,P. 1984. Activity patterns and time-budent in the goshawk Accipiter gentiles in a boreal forest area in Sweden. Ornis Fenn. 61:109～112.
117 陈华豪，张杰 1992 信息论在动物行为学中的应用。野生动物 (70) 3～5。
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86 Muller-Schwarze D 1968 Cirdadian rhythms of activity in Adelie penguin (Pygoscellis adeliae) during the austral summer[J]. Amarat Ros. Ser.,12:133～149.
87．张正旺，尹荣伦，郑光美，1989．笼养黄腹角雉繁殖期取食活动的研究[J]。动物学研究，10 (4)：331～339。
88 Bryant D M,Tatner P, 1988.Energetics of the annual cycle of dippers Cinclus cinclus[J].Ibis, 130:17～38.
89 Morton M L,1967.Diural feeding patterns in white-crowned sparrows Zonotrichia leucophrys gambelii[J].The Conder, 69:491～512.
90 Verbeek N A M, 1972.Daily andannual time budget of the yellow-billed magpie[J].Auk, 89:576～582.
91 杨晓君，文贤继，王淑珍，杨岚等，2000．笼养大紫胸鹦鹉取食活动。动物学研究 21(2)：115～120。
92 杨晓君，文贤继，杨岚，1994．笼养白腹锦鸡觅食活动的观察[J]。动物学研究，15(1)：49～54。
93 杨晓君，文贤继，杨岚，1995。笼养白腹锦鸡繁殖期的时间分配[J]。动物学研究，16 (2)：178～184。
94 杨晓君，周建平，杨岚，1991．棕胸竹鸡觅食行为的观察[J]。动物学研究，12(3)：259～264。
95 杨晓君，杨岚，1996。笼养绿孔雀行为活动时间分配的初步观察[J]。动物学报，42(增刊)：106～111。
96 田蕴，高玮，相桂权等，1996．人工饲养条件下花尾榛鸡的行为[J]。动物学报，42(增刊)：31～38。
97 杨晓君，杨岚，王淑珍，文贤继，1998．笼养大紫胸鹦鹉的活动时间分配[J]。动物学报，44 (3)：277～285。
98 Quinlan,E.E.&Baldassare,G.A. 1984. Activity budgets of nonbreeding green-winged teal on playa lakes in Texas. J.Wildl. Mgmt 48:838～845.
99 Bryant, C.J.& Thompson,D.B.A.1985.Gulls and Plovers: the ecology and behavior of mixed-species feeding groups. Beckenham:Croom Helm.
100 Carmi-Winkler, N.,Degen,A.A.&Pinshow, B. 1987. Seasonal time-energy budgets of free-livingchukars in the Negev desert. Condor 89:594～601.
101 Caraco,T. 1979. Time budgeting and group size: aTest of theory. Ecology 60:618～627.
102 East,M. 1980. Sex differences and the effect of temperature on the foraging behaviour of Robins Erithacus rubecula. Ibis 122:517～520.
103 East, M. 1982. Time-budgeting by European robins Erithacus rubecula: inter and intrasexual comparisons during autumn,winter and early spring. Ornis Scand 13:85～93.
104 Pearson O P, 1954.The daily energy requirements of a wild Anna Hummingbird. The Conder,56:317～322.
105 Hixon,M.A.,Carpenter, F.L.&Paton,D.C. 1983. Territory area,flower density, and time budgeting in hummingbirds:an experimental and theorectical anaslysis. Am.Nat. 122:366～391.
106 Gibb,J. 1956. Food,feeding habits and territory of the Rock Pipit Anthus spinoletta. Ibis 98: 506～530.
107 Hunter, M.L.,jr.,Witham,J.W.&Dow, H. 1984. Effects of a carbyl-induced depression in invertebrate abundance on the growth and behavior of American black duck and mallard ducklings. Can. J. Zool. 62:452～456.
108 Davies,N.B.&Lundberg,A. 1984. Food distribution and a variable mating system in the dunnock ,Prunella modularis. J.A nim. Ecol.53:895～912.
109 Enoksson,B.1990 Autumn territories and population regulation in the nuthatch Sitta europaea: An experimental study. J.Anim.
110 Ydenberg,R.C. 1984. The conflict between feeding and territorial defence in the great tit. Behav. Ecol. Sociobiol.15:103～108.
111 Davies,N.B.&Lundberg,A. 1985. The influence of food on time budgets and timing of breeding in the Dunnock Prunella modularis. Ibis 127:100～110.
112 Verner, J.1965. Time budget of the male long-billed marsh wren during the breeding season. Condor 67:125～139.
113 Maxson,S.J.&Oring,L.W. 1980. Breeding season time and energy budgets of the polyandrous spotted sandpiper. Behaviour 74:200～263.
114 Turner, A.K. 1982. Timing of laying by swallows (Hirundo rustica) and sand martins (Riparia riparia). J.Anim. Ecol. 51:29～46.
115 Siegfried,W.R. 1974. Time budgets of behavior among lesser scaups on Delta Marsh. J. Wildl. Mgmt 38:708～713.
116 Widen,P. 1984. Activity patterns and time-budent in the goshawk Accipiter gentiles in a boreal forest area in Sweden. Ornis Fenn. 61:109～112.
117 陈华豪，张杰 1992 信息论在动物行为学中的应用。野生动物 (70) 3～5。
118 郑光美 1995 鸟类学 北京师范大学出版社 (北京)：202．
119 江福霖等 1996 白鹳笼养繁殖的研究 中国动物园论文集 第3集 16～20 中国动物园协会编 中国大百科全书出版社上海分社出版
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