骨顶鸡的繁殖行为及其对人为干扰的响应
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摘要
2008年4月-11月,2009年4月-11月在黑龙江省安邦河自然保护区和大庆龙凤自然保护区对骨顶鸡的繁殖行为生态进行了研究,内容包括骨顶鸡的领域行为、种内巢寄生行为、生境选择行为以及骨顶鸡等水鸟对人为干扰的行为响应等。
研究结果显示骨顶鸡繁殖种群具有很强的领域性。领域在配对之后形成,繁殖对通过鸣叫、巡游以及建造“炫耀台”的方式昭示领域所有权,对同种和异种均有防御,领域行为在筑巢期领域活动最频繁,孵化期明显减弱,各阶段领域行为次数差异显著。雏鸟独立后领域性消失,重新集群。骨顶鸡的领域面积在1333 m2至5000 m2以上,全年5月至6月领域行为处于最高峰。越冬种群没有领域性。
骨顶鸡除了巢以外,繁殖季节会在领域内建造2-7个炫耀台,炫耀台形状上较巢简陋,且盖度低于巢,但炫耀台所在样方水深与巢下水深差异不显著。骨顶鸡在不同繁殖阶段对炫耀台和巢的利用率有所不同,筑巢期主要使用炫耀台用于理羽、休息、交尾、排泄以及储存食物等,而建巢之后以上大部分行为转移到巢上进行(除排泄外)。进入孵化期,对炫耀台的使用频率进一步降低。繁殖对每日在巢出现时间比例随繁殖阶段深入逐渐增加,在炫耀台每日出现的时间比例逐渐减少。此外,各个繁殖阶段雄性对炫耀台的利用率(每次出现时间、日使用次数、日使用时间)较雌鸟高,雌鸟对巢的利用率较高。推测炫耀台是骨顶鸡领域活动的产物,是一种领域昭示行为,同时炫耀台对于邻巢之间领域边界的划分与识别具有一定作用。
在2008-2009年两个繁殖季节对安邦河湿地和大庆龙凤湿地的骨顶鸡巢进行了调查,结果发现骨顶鸡具有比例很高的种内巢寄生现象,且骨顶鸡抵御种内巢寄生的方式类似于同属的美洲骨顶鸡(Fulilca americana):通过埋卵、逐出和啄破拒绝寄生卵,结合调查中发现的骨顶鸡的领域行为以及在领域内筑多个炫耀台(display platform)等现象对骨顶鸡抵御种内巢寄生的机制进行了探讨。
领域行为及其建造炫耀台作为抵御种内巢寄生的策略可以较好的解释领域性产生的机制,这种繁殖策略选择有利于增加其适合度,使种群达到稳定状态。调查发现,除同种其他个体之外,骨顶鸡对红头潜鸭的驱逐次数最多,红头潜鸭也是唯一进入骨顶鸡“炫耀台”内的异种,根据红头潜鸭具有巢寄生现象以及其他骨顶属物种有被鸭科鸟类寄生的记录,推测骨顶鸡领域行为产生机制可能与巢寄生有关。
采用主成分分析和逐步判别法,结合观察法对安邦河湿地和大庆龙凤湿地骨顶鸡的巢址偏好特征进行了研究。主成分分析和逐步判别结果表明：(1)骨顶鸡选择盖度适中、高度适中、距明水面较近、水深适中、较为稀疏的芦苇丛和蒲草丛中,(2)人为干扰因素对骨顶鸡的巢址选择影响较小,而种内关系及种间关系对其巢址选择影响较大,(3)骨顶鸡繁殖期的领域性对其生境选择具有重要影响。
笔者分别于2008年4月至6月和2009年4月至6月,采用目标动物取样法对黑龙江省安邦河湿地和大庆龙凤湿地人为干扰对骨顶鸡等水鸟的影响进行了研究。结果表明：骨顶鸡对行人和机动车辆两种干扰的反应距离类似,但耐受程度有所不同,骨顶鸡对机动车辆干扰的耐受距离要大于对游客等行人干扰的耐受距离,说明骨顶鸡对于噪声干扰有所回避。在以游客等行人干扰为主的安邦河湿地,骨顶鸡以及其他游禽对干扰产生反应的比例要远高于以机动车辆干扰为主的龙凤湿地,说明相对于人产生的干扰骨顶鸡对机动车辆更容易耐受；在安邦河湿地,骨顶鸡对划船的反应距离以及反应程度远高于行人行走,说明骨顶鸡对于侵入性的干扰更为敏感。
采用固定样线法和样点法对安邦河湿地干扰区(旅游区)和核心区骨顶鸡的数量监测结果显示：干扰区骨顶鸡的数量不少于核心区,且个别月份(10月)显著高于核心区；对其他水禽的种类和数量调查也显示安邦河湿地人为干扰对水鸟的分布及数量影响较小,仅个别敏感种类在旅游区缺乏观测记录。相对于旅游活动产生的人为干扰,骨顶鸡对人为造成的水位变动更为敏感,繁殖种群和越冬种群的分布均受水位变动的影响。
尽管人为干扰因素对骨顶鸡的数量、分布以及巢址选择没有显著影响,但其很可能与生境质量一起对骨顶鸡的繁殖时间产生影响。靠近岸边(干扰强烈、生境质量较差)的繁殖对繁殖时间较晚。
基于上述结果,对骨顶鸡繁殖策略得出以下结论：
(1)骨顶鸡产卵之前领域活动的能量投入较高,建立领域具有占有资源、监护配偶以及抵御巢寄生的作用,随着繁殖阶段的深入,与邻巢之间倾向于选择能量消耗较小的领域行为类型,能量投入转向繁殖后代。
(2)骨顶鸡存在较高比例的种内巢寄生现象,而种内巢寄生对宿主而言是耗费成本的,因此骨顶鸡进化出卵识别能力和拒卵行为,并通过领域监护以及建筑炫耀台的方式减少被寄生的可能。
(3)骨顶鸡巢址选择倾向于生境质量较高的区域,有利于繁殖产出,但领域性及资源可获得性会影响骨顶鸡的巢址选择,种内竞争使得部分繁殖对无法获得高质量的生境,但这些繁殖对可能会通过延迟繁殖时间的方式等待生境条件改善,从而提高繁殖产出。
(4)骨顶鸡对人为干扰的行为反应较鸭类等其他水鸟要少,且程度较低,领域性使边缘个体受到的干扰较多,但边缘个体会以耐受的方式减少惊飞、惊跑造成的能量消耗,更多选择游向隐蔽处这样一种回避干扰的行为方式。
A scientific research on behavioral ecology of Common Coot was made in Anbanghe Nature Reserve and Daqing Longfeng Nature Reserve since April to November of 2008 and 2009. Research contents included territory behavior, conspecific brood parasitism, habitat selection, as well as coots' behavioral response to human disturbance.
Territory and territorial behavior of Common Coot were observed in both sites in breeding season, territory were formed after paired, and both sexes display the rights and protected it through calls, patrolling and "display platform" construction. Both individuals of coots and other species were drove out when they approached the boundary. Territory activities became severe since May to June, the territorility level was highest in the nest building period and weaker since then. As territory owners engage in territorial fights and contests of their neighbors, and avoid the potential costs of escalated fights was reasonable to interprate the weaker of territory activities; more energy were turned into breeding after egg laying was the other reason for interpreting the weaker of territory activities. After the young was independent, territory behaviors disappeared, and the coots gathered again. Territory size differed from 1 333 m2 to above 5 000 m2. Wintering population was observed in Poyang Lake of Jiangxi province; the coots gathered in the open water and did not present territory behaviors in wintering sites.
Similar to the American coot (Fulica americana), each pair of Common Coot, build 2-7 display platforms in their territory during the breeding season, besides the nest. The functionality of display platforms of the coots and figure differences of them with nests has been discussed; nests were more refined and significantly higher than the platforms. Display platforms were used primarily for roosting, preening, and mating in reproduction periods, and the use ration of platforms decreased with the development of reproduction stages, contrarily, time spend on nests increased. Besides these, the platforms might play an important role in confirming and recognizing the territory boundaries. Besides these, for the platforms, use ration (time of every present, day present times and day total present time) of the male was higher than the female in different breeding stages, for the nest, the result was contrary. Platforms construction was most active in nest building period, and weaker in next stages. However, the coots would add new platforms near conflict points when they were incubation. It seems that the construction of platforms was a result of neighbor conflicts, so I inferred the platforms were a kind of territory display, and they might be used as a reference of territory boundary reorganization.
Conspecific brood parasitism of Common Coot was observed in both study sites. Similar as American Coot, egg rejection was used for parasitism defence; parasites' eggs were buried, driven out and pecked by the hosts. Besides these manners, territory behaviors and the construction of display platforms might play an important role in defending parasitism In species driven by the coots, Common Parchard was observed most, and they occupied the coots' structures, as inter-specific nest parasitism of Common Parchard had been reported, and ducks were detected in the list of parasites of other coots, so I inferred the severe territory behaviors of coots to Common Parchard might be caused by brood parasitism.
Effects of human activity on the behavior of waterbirds were checked using focal-animal sampling method in two study sites from April to June of 2008 and 2009. The results showed that the responding distances of the Common Coots to human walking and motor vehicles were similar, however, the tolerance degrees were different, tolerance distance of the coots to motor vehicles were larger than human walking, and the percentage of responses to disturbance was higher in Anbanghe wetland where the main disturbance type was tourists' walking, which indicated that the coots and other swimming birds were more tolerant to motor vehicles compared with walking. In Anbanghe wetland, the responding distance of the coots to boating disturbance was much larger than walking; it means the coots were less tolerant to invading activities.
Fixed line transect method and sample plot method were used combined to investigate the amount and distribution of coots and other species in disturbance area (tourist area) and core area of Anbanghe wetland. Results indicated that the amount of coots in disturbance area were similar as the core area and even higher in disturbance area in October as more coots gathered in the open water of the lake in the disturbance area. Water depth and open water area were more important to coots compared with recreation disturbance. The species abundance and distribution were not significant influenced by recreation disturbance in Anbanghe wetland.
Though the coots shows little response to human disturbance on behaviors, amount and distribution, human disturbance combined with habitat quality might delay the breeding date of the coots.
Principal component analysis and stepwise discriminate analysis were used associated with observation to investigate the nest-site selection of coots in breeding season. Results showed that:(1) coots favored to nest in area with middle vegetation coverage and height, moderate water level, near to open water and thin reeds and cattails; (2) impact of human disturbance on nest site selection was weaker, intra and inter species attractions and competition were more influential factors for coots' nest site selection. (3) territoriality played important roles in habitat selection of the coots.
On the basis of the aforesaid results, the coots adopted breeding tactics as followed:
(1) More energy were paid on territory activities before egg laying, and territory was important in occupying resources, mate-guarding and defending parasitism; after egg laying, more energy was turned into breeding offsprings, neighbors tended to adopt behaviors less energy cost in territory activities.
(2) High proportion of conspecific brood parasitism existed in coots, and parasitism was cost for the hosts as fitness reduction, so defense mechanism was important for the species with CBP, besides egg rejection based on egg recognization, the coots adopted territory guarding and platforms construction to reduce the parasite opportunities by other females.
(3) Coots tend to chose high quality habitat for nesting, which wound be advantages for reproduction, however, territory competition and resource availability will made some pairs unable to obtain these areas, they might chose to delay the breeding date until habitat changes better.
(4) Coots response to human disturbance less sensitively compared with ducks, and avoidance degree was lower, pairs distributed in the edge were disturbed more frequently than the central ones, they adopted less energy cost response to the disturbance, such as swim away which was more used by the edge pairs compared with flush and run.
研究结果显示骨顶鸡繁殖种群具有很强的领域性。领域在配对之后形成,繁殖对通过鸣叫、巡游以及建造“炫耀台”的方式昭示领域所有权,对同种和异种均有防御,领域行为在筑巢期领域活动最频繁,孵化期明显减弱,各阶段领域行为次数差异显著。雏鸟独立后领域性消失,重新集群。骨顶鸡的领域面积在1333 m2至5000 m2以上,全年5月至6月领域行为处于最高峰。越冬种群没有领域性。
骨顶鸡除了巢以外,繁殖季节会在领域内建造2-7个炫耀台,炫耀台形状上较巢简陋,且盖度低于巢,但炫耀台所在样方水深与巢下水深差异不显著。骨顶鸡在不同繁殖阶段对炫耀台和巢的利用率有所不同,筑巢期主要使用炫耀台用于理羽、休息、交尾、排泄以及储存食物等,而建巢之后以上大部分行为转移到巢上进行(除排泄外)。进入孵化期,对炫耀台的使用频率进一步降低。繁殖对每日在巢出现时间比例随繁殖阶段深入逐渐增加,在炫耀台每日出现的时间比例逐渐减少。此外,各个繁殖阶段雄性对炫耀台的利用率(每次出现时间、日使用次数、日使用时间)较雌鸟高,雌鸟对巢的利用率较高。推测炫耀台是骨顶鸡领域活动的产物,是一种领域昭示行为,同时炫耀台对于邻巢之间领域边界的划分与识别具有一定作用。
在2008-2009年两个繁殖季节对安邦河湿地和大庆龙凤湿地的骨顶鸡巢进行了调查,结果发现骨顶鸡具有比例很高的种内巢寄生现象,且骨顶鸡抵御种内巢寄生的方式类似于同属的美洲骨顶鸡(Fulilca americana):通过埋卵、逐出和啄破拒绝寄生卵,结合调查中发现的骨顶鸡的领域行为以及在领域内筑多个炫耀台(display platform)等现象对骨顶鸡抵御种内巢寄生的机制进行了探讨。
领域行为及其建造炫耀台作为抵御种内巢寄生的策略可以较好的解释领域性产生的机制,这种繁殖策略选择有利于增加其适合度,使种群达到稳定状态。调查发现,除同种其他个体之外,骨顶鸡对红头潜鸭的驱逐次数最多,红头潜鸭也是唯一进入骨顶鸡“炫耀台”内的异种,根据红头潜鸭具有巢寄生现象以及其他骨顶属物种有被鸭科鸟类寄生的记录,推测骨顶鸡领域行为产生机制可能与巢寄生有关。
采用主成分分析和逐步判别法,结合观察法对安邦河湿地和大庆龙凤湿地骨顶鸡的巢址偏好特征进行了研究。主成分分析和逐步判别结果表明：(1)骨顶鸡选择盖度适中、高度适中、距明水面较近、水深适中、较为稀疏的芦苇丛和蒲草丛中,(2)人为干扰因素对骨顶鸡的巢址选择影响较小,而种内关系及种间关系对其巢址选择影响较大,(3)骨顶鸡繁殖期的领域性对其生境选择具有重要影响。
笔者分别于2008年4月至6月和2009年4月至6月,采用目标动物取样法对黑龙江省安邦河湿地和大庆龙凤湿地人为干扰对骨顶鸡等水鸟的影响进行了研究。结果表明：骨顶鸡对行人和机动车辆两种干扰的反应距离类似,但耐受程度有所不同,骨顶鸡对机动车辆干扰的耐受距离要大于对游客等行人干扰的耐受距离,说明骨顶鸡对于噪声干扰有所回避。在以游客等行人干扰为主的安邦河湿地,骨顶鸡以及其他游禽对干扰产生反应的比例要远高于以机动车辆干扰为主的龙凤湿地,说明相对于人产生的干扰骨顶鸡对机动车辆更容易耐受；在安邦河湿地,骨顶鸡对划船的反应距离以及反应程度远高于行人行走,说明骨顶鸡对于侵入性的干扰更为敏感。
采用固定样线法和样点法对安邦河湿地干扰区(旅游区)和核心区骨顶鸡的数量监测结果显示：干扰区骨顶鸡的数量不少于核心区,且个别月份(10月)显著高于核心区；对其他水禽的种类和数量调查也显示安邦河湿地人为干扰对水鸟的分布及数量影响较小,仅个别敏感种类在旅游区缺乏观测记录。相对于旅游活动产生的人为干扰,骨顶鸡对人为造成的水位变动更为敏感,繁殖种群和越冬种群的分布均受水位变动的影响。
尽管人为干扰因素对骨顶鸡的数量、分布以及巢址选择没有显著影响,但其很可能与生境质量一起对骨顶鸡的繁殖时间产生影响。靠近岸边(干扰强烈、生境质量较差)的繁殖对繁殖时间较晚。
基于上述结果,对骨顶鸡繁殖策略得出以下结论：
(1)骨顶鸡产卵之前领域活动的能量投入较高,建立领域具有占有资源、监护配偶以及抵御巢寄生的作用,随着繁殖阶段的深入,与邻巢之间倾向于选择能量消耗较小的领域行为类型,能量投入转向繁殖后代。
(2)骨顶鸡存在较高比例的种内巢寄生现象,而种内巢寄生对宿主而言是耗费成本的,因此骨顶鸡进化出卵识别能力和拒卵行为,并通过领域监护以及建筑炫耀台的方式减少被寄生的可能。
(3)骨顶鸡巢址选择倾向于生境质量较高的区域,有利于繁殖产出,但领域性及资源可获得性会影响骨顶鸡的巢址选择,种内竞争使得部分繁殖对无法获得高质量的生境,但这些繁殖对可能会通过延迟繁殖时间的方式等待生境条件改善,从而提高繁殖产出。
(4)骨顶鸡对人为干扰的行为反应较鸭类等其他水鸟要少,且程度较低,领域性使边缘个体受到的干扰较多,但边缘个体会以耐受的方式减少惊飞、惊跑造成的能量消耗,更多选择游向隐蔽处这样一种回避干扰的行为方式。
A scientific research on behavioral ecology of Common Coot was made in Anbanghe Nature Reserve and Daqing Longfeng Nature Reserve since April to November of 2008 and 2009. Research contents included territory behavior, conspecific brood parasitism, habitat selection, as well as coots' behavioral response to human disturbance.
Territory and territorial behavior of Common Coot were observed in both sites in breeding season, territory were formed after paired, and both sexes display the rights and protected it through calls, patrolling and "display platform" construction. Both individuals of coots and other species were drove out when they approached the boundary. Territory activities became severe since May to June, the territorility level was highest in the nest building period and weaker since then. As territory owners engage in territorial fights and contests of their neighbors, and avoid the potential costs of escalated fights was reasonable to interprate the weaker of territory activities; more energy were turned into breeding after egg laying was the other reason for interpreting the weaker of territory activities. After the young was independent, territory behaviors disappeared, and the coots gathered again. Territory size differed from 1 333 m2 to above 5 000 m2. Wintering population was observed in Poyang Lake of Jiangxi province; the coots gathered in the open water and did not present territory behaviors in wintering sites.
Similar to the American coot (Fulica americana), each pair of Common Coot, build 2-7 display platforms in their territory during the breeding season, besides the nest. The functionality of display platforms of the coots and figure differences of them with nests has been discussed; nests were more refined and significantly higher than the platforms. Display platforms were used primarily for roosting, preening, and mating in reproduction periods, and the use ration of platforms decreased with the development of reproduction stages, contrarily, time spend on nests increased. Besides these, the platforms might play an important role in confirming and recognizing the territory boundaries. Besides these, for the platforms, use ration (time of every present, day present times and day total present time) of the male was higher than the female in different breeding stages, for the nest, the result was contrary. Platforms construction was most active in nest building period, and weaker in next stages. However, the coots would add new platforms near conflict points when they were incubation. It seems that the construction of platforms was a result of neighbor conflicts, so I inferred the platforms were a kind of territory display, and they might be used as a reference of territory boundary reorganization.
Conspecific brood parasitism of Common Coot was observed in both study sites. Similar as American Coot, egg rejection was used for parasitism defence; parasites' eggs were buried, driven out and pecked by the hosts. Besides these manners, territory behaviors and the construction of display platforms might play an important role in defending parasitism In species driven by the coots, Common Parchard was observed most, and they occupied the coots' structures, as inter-specific nest parasitism of Common Parchard had been reported, and ducks were detected in the list of parasites of other coots, so I inferred the severe territory behaviors of coots to Common Parchard might be caused by brood parasitism.
Effects of human activity on the behavior of waterbirds were checked using focal-animal sampling method in two study sites from April to June of 2008 and 2009. The results showed that the responding distances of the Common Coots to human walking and motor vehicles were similar, however, the tolerance degrees were different, tolerance distance of the coots to motor vehicles were larger than human walking, and the percentage of responses to disturbance was higher in Anbanghe wetland where the main disturbance type was tourists' walking, which indicated that the coots and other swimming birds were more tolerant to motor vehicles compared with walking. In Anbanghe wetland, the responding distance of the coots to boating disturbance was much larger than walking; it means the coots were less tolerant to invading activities.
Fixed line transect method and sample plot method were used combined to investigate the amount and distribution of coots and other species in disturbance area (tourist area) and core area of Anbanghe wetland. Results indicated that the amount of coots in disturbance area were similar as the core area and even higher in disturbance area in October as more coots gathered in the open water of the lake in the disturbance area. Water depth and open water area were more important to coots compared with recreation disturbance. The species abundance and distribution were not significant influenced by recreation disturbance in Anbanghe wetland.
Though the coots shows little response to human disturbance on behaviors, amount and distribution, human disturbance combined with habitat quality might delay the breeding date of the coots.
Principal component analysis and stepwise discriminate analysis were used associated with observation to investigate the nest-site selection of coots in breeding season. Results showed that:(1) coots favored to nest in area with middle vegetation coverage and height, moderate water level, near to open water and thin reeds and cattails; (2) impact of human disturbance on nest site selection was weaker, intra and inter species attractions and competition were more influential factors for coots' nest site selection. (3) territoriality played important roles in habitat selection of the coots.
On the basis of the aforesaid results, the coots adopted breeding tactics as followed:
(1) More energy were paid on territory activities before egg laying, and territory was important in occupying resources, mate-guarding and defending parasitism; after egg laying, more energy was turned into breeding offsprings, neighbors tended to adopt behaviors less energy cost in territory activities.
(2) High proportion of conspecific brood parasitism existed in coots, and parasitism was cost for the hosts as fitness reduction, so defense mechanism was important for the species with CBP, besides egg rejection based on egg recognization, the coots adopted territory guarding and platforms construction to reduce the parasite opportunities by other females.
(3) Coots tend to chose high quality habitat for nesting, which wound be advantages for reproduction, however, territory competition and resource availability will made some pairs unable to obtain these areas, they might chose to delay the breeding date until habitat changes better.
(4) Coots response to human disturbance less sensitively compared with ducks, and avoidance degree was lower, pairs distributed in the edge were disturbed more frequently than the central ones, they adopted less energy cost response to the disturbance, such as swim away which was more used by the edge pairs compared with flush and run.
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