食物对越冬小白额雁分布、能量与氮平衡以及行为的影响
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摘要
全球受胁物种小白额雁在近几十年来经历了快速的数量下降和分布缩减。而在全球最大的越冬地,长江中下游湿地的小白额雁现状及历史变化仍无全面理解。同时,人们始终未能解释为何越冬豆雁和白额雁在长江中下游湿地广泛分布、而小白额雁高度集中分布在东洞庭湖。此外,越冬小白额雁的觅食行为,一直未被研究。本论文围绕以上几个核心问题进行了研究,并取得下列结果：
1.我国越冬小白额雁数量和分布的变化
近几十年来,全球小白额雁经历了迅速的数量下降和分布缩减。本研究之前的全球种群数量估计为25000至28000只,其中约20000只在我国东洞庭湖越冬。本研究通过整合、分析已发表和未发表数据,首次掌握了我国越冬小白额雁的数量、分布及其历史变化。本研究发现,经历了最近二十余年的数量减少与分布缩减,我国小白额雁目前主要集中在东洞庭湖几个越冬位点。其稳定的数量表明小白额雁并非极度受胁,但高度集中的分布表明该物种极易受到栖息地破坏等因素的影响。因此,对该物种的有效保护,迫切需要了解形成其独特分布的原因。
2.不同类型觅食地小白额雁的能量平衡
在众多影响物种分布的因素当中,食物最可能决定越冬雁类分布。本研究利用植物中不可消化的内源标记物,研究了整个越冬期不同类型觅食地上小白额雁的能量平衡。结果表明,在看麦娘、狗牙根及江南荸荠优势的斑块化栖息地觅食的小白额雁,在整个越冬期可以维持总体上的正能量平衡；而在苔草优势的栖息地上,小白额雁在整个越冬期都无法维持正能量平衡。而相较于看麦娘、狗牙根及江南荸荠栖息地斑块化的分布,苔草滩涂在长江中下游湿地均广泛分布。本研究的结果解释了为何小白额雁没有分布在苔草优势的湿地。但同时,本研究提出了两个新问题：既然没有能量收益,小白额雁为何取食苔草?长江中下游湿地越冬白额雁和豆雁的广泛分布,能否也从相同的角度来解释?
3.不同类型觅食地小白额雁的氮平衡
当主要食物缺乏某种必需元素时,动物可能会切换到另一种食物来补充该必需元素,从而使其呈现异常分布。对雁类来说,这种必需元素通常为氮。本研究利用植物中不可消化的内源标记物,计算了两种觅食地上小白额雁的氮平衡。结果显示,在整个越冬期,能够为小白额雁带来正能量平衡的食物也能够带来正的氮平衡,而不能提供足够能量摄入的食物,亦不能提供足够的氮摄入。因此,小白额雁取食苔草并非为了摄入氮元素,而可能是为了避免人为干扰而采取的权宜策略。
4.升金湖取食苔草的白额雁与豆雁的越冬期能量平衡
如果取食苔草时的低能量摄入能够解释小白额雁为何不分布在在苔草优势的湿地,那么广泛分布在长江中下游苔草湿地的白额雁和豆雁应当可以通过取食苔草而获得正能量平衡。本研究估算了利用升金湖苔草滩的越冬白额雁及豆雁的食物组成和能量平衡。结果表明,两种雁均主要取食苔草,而且可以在整个越冬期维持总体上的正能量平衡。该结果解释了白额雁及豆雁在长江流域的广泛分布,加强了关于小白额雁分布影响因素的结论,并验证了“与小体型动物相比,大体型动物可以耐受低食物质量,但需要更高的食物供应量”这一觅食理论中的经典观点。
5.小白额雁在不同觅食地的日活动时间分配
大量研究表明,高觅食时间投入通常伴随了低食物可利用度或低能量摄入。然而觅食时间投入受到许多因素影响,因此高觅食时间投入未必一定伴随低能量摄入,但目前尚无研究报道这一现象。本研究首次观察并比较了分别在大西湖和采桑湖觅食的小白额雁的日活动时间分配。结果表明,与在低能量平衡的大西湖相比,在高能量平衡的采桑湖,小白额雁的觅食比例更高,休息比例更低。而且在本研究中,其它影响觅食时间的因素(如种内、种间竞争,人类干扰、觅食者掌握的觅食地信息及生活史阶段)不存在或影响较小。该研究表明,仅通过日活动时间分配(如觅食时间和比例)不足以推断动物的能量平衡。
以上五项研究,首次综述了我国小白额雁的历史与现状,从能量与氮代谢角度阐明了小白额雁及邻域分布物种白额雁与豆雁的分布形成机制,首次描述了在我国越冬小白额雁在整个越冬期的日活动时间分配,并发现了能量平衡与日活动时间分配之间的异常关系。
The globally threatened Lesser White-fronted Goose Anser erythropus underwent a rapid decline in abundance and contraction in distribution. Little is known about the status and changes of this species in Yangtze wetlands, the biggest wintering ground in the world. Another unsolved question about this species is its extraordinarily localized distribution, comparing with other species such as the Bean Goose Anser fabalis serrirostris and Greater White-fronted Goose Anser albifrons which widespread within Yangtze River wetlands. In addition, the foraging behavior of wintering Lesser White-fronted Geese is poorly understood. In order to answer these questions, I conducted five studies in this thesis:
1. Changes in distribution and abundance of wintering Lesser White-fronted Geese in eastern China
The global population of Lesser White-fronted Geese has decreased rapidly in recent decades. Before this study, the world population estimate of this population was25000to28000, among which c.20000individuals wintered in East Dongting Lake, China. By analyzing published and unpublished data, we present the first distribution, abundance and population changes in recent two decades. After a rapid historical population decline, the geese now highly concentrate at a few sites within East Dongting Lake. The relatively stable numbers at these sites suggest the population is not currently threatened, but the localized distribution makes the species vulnerable. In order for effective conservation, factors contributing to the unique distribution of this species need to be investigated.
2. Energy budgets of Lesser White-fronted Geese at different habitat types
Among factors influencing distribution, food is most likely to determine the distribution of wintering geese. By using the endogenous indigestible marker in plants, we estimated the energy budgets of Lesser White-fronted Geese foraging on two types of habitats. The results show the Alopecurus, Cynodon and Eleocharis, patchily distributed within East Dongting Lake, could provide the geese with overall positive energy budgets through the entire wintering period, while geese failed to maintain the energetic balance by feeding on Carex, the dominant species along Yangtze River wetlands. Our finding explains why Lesser White-fronted Geese are not widely distributed. However, this study also sets two new questions:why the geese foraged on energy-poor Carex, and whether the wide distribution of Greater White-fronted Geese and Bean Geese can be explained from the same perspective.
3. Nitrogen budgets of Lesser White-fronted Geese at different habitat types
When the main diet is short of some necessary element (for herbivorous geese this element is often nitrogen), consumers may temporarily shift to a rare diet to supplement this nutrient. This may explain species abnormal distributions. Under this hypothesis, we examined the nitrogen budgets of Lesser White-fronted Geese foraging on energy-rich and-poor diets, by using the endogenous indigestible marker in plants. The results show that the energy-rich diet could provide the geese with positive nitrogen budget all through the wintering period, but the energy-poor diet failed to support the geese with nitrogen balance. In conclusion, Lesser White-fronted Geese did not feed on Carex for nitrogen intake, but probably did so as a result of the anthropogenic disturbance.
4. The energy budgets of wintering Greater White-fronted Geese and Bean Geese feeding on Carex at Shengjin Lake
If the localized distribution pattern of Lesser White-fronted Geese could be explained by its failure by grazing Carex, the widespread Greater White-fronted Geese and Bean Geese through Yangtze River wetlands should also result from the benefit from grazing Carex. We estimated energy budgets of these two species foraging on Carex at Shengjin Lake, through the entire wintering period. Our results show that both species could gain overall positive energy budgets by grazing Carex through the wintering period. This result explains the wide distribution of both species.
5. The unexpected time-activity budgets of Lesser White-fronted Geese on two habitat types
Elevated investment in foraging time is usually associated with low food availability or low metabolizable energy intake. But this may not always be the case. Comparing full-day activity budgets through the course of the winter, we found an unexpected difference in activity budgets of wintering Lesser White-fronted Geese Anser erythropus at two sites in close proximity, which differed in vegetation, within East Dongting Lake, China. Counter to expectations, geese fed for significantly more of the time and spent significantly less time resting at the site which provided energy-rich food than the energy-poor one. Given the little time invested in vigilance and aggression, and the absence of other factors influencing activity budgets, we argue that the unexpected differences in investment in feeding activity were primarily shaped by the high quality but low quantity of the food at one site (causing a constraint intake rates that elevated harvest time) and high quantity low quality at the other, where the lower investment in feeding resulted from limits on digestion rate. Our finding urges caution regarding drawing conclusions on energy budgets and food preferences based on activity budgets alone.
The five studies provide the first overview of the status and history of Lesser White-fronted Geese, explain the distribution of the Lesser White-fronted Goose and its neighboring Greater White-fronted Geese and Bean Geese from the energetic and nutrient perspectives, find the effect of diet type on behavior and present the first time-activity budgets of Lesser White-fronted Geese on natural habitats during the entire wintering period.
1.我国越冬小白额雁数量和分布的变化
近几十年来,全球小白额雁经历了迅速的数量下降和分布缩减。本研究之前的全球种群数量估计为25000至28000只,其中约20000只在我国东洞庭湖越冬。本研究通过整合、分析已发表和未发表数据,首次掌握了我国越冬小白额雁的数量、分布及其历史变化。本研究发现,经历了最近二十余年的数量减少与分布缩减,我国小白额雁目前主要集中在东洞庭湖几个越冬位点。其稳定的数量表明小白额雁并非极度受胁,但高度集中的分布表明该物种极易受到栖息地破坏等因素的影响。因此,对该物种的有效保护,迫切需要了解形成其独特分布的原因。
2.不同类型觅食地小白额雁的能量平衡
在众多影响物种分布的因素当中,食物最可能决定越冬雁类分布。本研究利用植物中不可消化的内源标记物,研究了整个越冬期不同类型觅食地上小白额雁的能量平衡。结果表明,在看麦娘、狗牙根及江南荸荠优势的斑块化栖息地觅食的小白额雁,在整个越冬期可以维持总体上的正能量平衡；而在苔草优势的栖息地上,小白额雁在整个越冬期都无法维持正能量平衡。而相较于看麦娘、狗牙根及江南荸荠栖息地斑块化的分布,苔草滩涂在长江中下游湿地均广泛分布。本研究的结果解释了为何小白额雁没有分布在苔草优势的湿地。但同时,本研究提出了两个新问题：既然没有能量收益,小白额雁为何取食苔草?长江中下游湿地越冬白额雁和豆雁的广泛分布,能否也从相同的角度来解释?
3.不同类型觅食地小白额雁的氮平衡
当主要食物缺乏某种必需元素时,动物可能会切换到另一种食物来补充该必需元素,从而使其呈现异常分布。对雁类来说,这种必需元素通常为氮。本研究利用植物中不可消化的内源标记物,计算了两种觅食地上小白额雁的氮平衡。结果显示,在整个越冬期,能够为小白额雁带来正能量平衡的食物也能够带来正的氮平衡,而不能提供足够能量摄入的食物,亦不能提供足够的氮摄入。因此,小白额雁取食苔草并非为了摄入氮元素,而可能是为了避免人为干扰而采取的权宜策略。
4.升金湖取食苔草的白额雁与豆雁的越冬期能量平衡
如果取食苔草时的低能量摄入能够解释小白额雁为何不分布在在苔草优势的湿地,那么广泛分布在长江中下游苔草湿地的白额雁和豆雁应当可以通过取食苔草而获得正能量平衡。本研究估算了利用升金湖苔草滩的越冬白额雁及豆雁的食物组成和能量平衡。结果表明,两种雁均主要取食苔草,而且可以在整个越冬期维持总体上的正能量平衡。该结果解释了白额雁及豆雁在长江流域的广泛分布,加强了关于小白额雁分布影响因素的结论,并验证了“与小体型动物相比,大体型动物可以耐受低食物质量,但需要更高的食物供应量”这一觅食理论中的经典观点。
5.小白额雁在不同觅食地的日活动时间分配
大量研究表明,高觅食时间投入通常伴随了低食物可利用度或低能量摄入。然而觅食时间投入受到许多因素影响,因此高觅食时间投入未必一定伴随低能量摄入,但目前尚无研究报道这一现象。本研究首次观察并比较了分别在大西湖和采桑湖觅食的小白额雁的日活动时间分配。结果表明,与在低能量平衡的大西湖相比,在高能量平衡的采桑湖,小白额雁的觅食比例更高,休息比例更低。而且在本研究中,其它影响觅食时间的因素(如种内、种间竞争,人类干扰、觅食者掌握的觅食地信息及生活史阶段)不存在或影响较小。该研究表明,仅通过日活动时间分配(如觅食时间和比例)不足以推断动物的能量平衡。
以上五项研究,首次综述了我国小白额雁的历史与现状,从能量与氮代谢角度阐明了小白额雁及邻域分布物种白额雁与豆雁的分布形成机制,首次描述了在我国越冬小白额雁在整个越冬期的日活动时间分配,并发现了能量平衡与日活动时间分配之间的异常关系。
The globally threatened Lesser White-fronted Goose Anser erythropus underwent a rapid decline in abundance and contraction in distribution. Little is known about the status and changes of this species in Yangtze wetlands, the biggest wintering ground in the world. Another unsolved question about this species is its extraordinarily localized distribution, comparing with other species such as the Bean Goose Anser fabalis serrirostris and Greater White-fronted Goose Anser albifrons which widespread within Yangtze River wetlands. In addition, the foraging behavior of wintering Lesser White-fronted Geese is poorly understood. In order to answer these questions, I conducted five studies in this thesis:
1. Changes in distribution and abundance of wintering Lesser White-fronted Geese in eastern China
The global population of Lesser White-fronted Geese has decreased rapidly in recent decades. Before this study, the world population estimate of this population was25000to28000, among which c.20000individuals wintered in East Dongting Lake, China. By analyzing published and unpublished data, we present the first distribution, abundance and population changes in recent two decades. After a rapid historical population decline, the geese now highly concentrate at a few sites within East Dongting Lake. The relatively stable numbers at these sites suggest the population is not currently threatened, but the localized distribution makes the species vulnerable. In order for effective conservation, factors contributing to the unique distribution of this species need to be investigated.
2. Energy budgets of Lesser White-fronted Geese at different habitat types
Among factors influencing distribution, food is most likely to determine the distribution of wintering geese. By using the endogenous indigestible marker in plants, we estimated the energy budgets of Lesser White-fronted Geese foraging on two types of habitats. The results show the Alopecurus, Cynodon and Eleocharis, patchily distributed within East Dongting Lake, could provide the geese with overall positive energy budgets through the entire wintering period, while geese failed to maintain the energetic balance by feeding on Carex, the dominant species along Yangtze River wetlands. Our finding explains why Lesser White-fronted Geese are not widely distributed. However, this study also sets two new questions:why the geese foraged on energy-poor Carex, and whether the wide distribution of Greater White-fronted Geese and Bean Geese can be explained from the same perspective.
3. Nitrogen budgets of Lesser White-fronted Geese at different habitat types
When the main diet is short of some necessary element (for herbivorous geese this element is often nitrogen), consumers may temporarily shift to a rare diet to supplement this nutrient. This may explain species abnormal distributions. Under this hypothesis, we examined the nitrogen budgets of Lesser White-fronted Geese foraging on energy-rich and-poor diets, by using the endogenous indigestible marker in plants. The results show that the energy-rich diet could provide the geese with positive nitrogen budget all through the wintering period, but the energy-poor diet failed to support the geese with nitrogen balance. In conclusion, Lesser White-fronted Geese did not feed on Carex for nitrogen intake, but probably did so as a result of the anthropogenic disturbance.
4. The energy budgets of wintering Greater White-fronted Geese and Bean Geese feeding on Carex at Shengjin Lake
If the localized distribution pattern of Lesser White-fronted Geese could be explained by its failure by grazing Carex, the widespread Greater White-fronted Geese and Bean Geese through Yangtze River wetlands should also result from the benefit from grazing Carex. We estimated energy budgets of these two species foraging on Carex at Shengjin Lake, through the entire wintering period. Our results show that both species could gain overall positive energy budgets by grazing Carex through the wintering period. This result explains the wide distribution of both species.
5. The unexpected time-activity budgets of Lesser White-fronted Geese on two habitat types
Elevated investment in foraging time is usually associated with low food availability or low metabolizable energy intake. But this may not always be the case. Comparing full-day activity budgets through the course of the winter, we found an unexpected difference in activity budgets of wintering Lesser White-fronted Geese Anser erythropus at two sites in close proximity, which differed in vegetation, within East Dongting Lake, China. Counter to expectations, geese fed for significantly more of the time and spent significantly less time resting at the site which provided energy-rich food than the energy-poor one. Given the little time invested in vigilance and aggression, and the absence of other factors influencing activity budgets, we argue that the unexpected differences in investment in feeding activity were primarily shaped by the high quality but low quantity of the food at one site (causing a constraint intake rates that elevated harvest time) and high quantity low quality at the other, where the lower investment in feeding resulted from limits on digestion rate. Our finding urges caution regarding drawing conclusions on energy budgets and food preferences based on activity budgets alone.
The five studies provide the first overview of the status and history of Lesser White-fronted Geese, explain the distribution of the Lesser White-fronted Goose and its neighboring Greater White-fronted Geese and Bean Geese from the energetic and nutrient perspectives, find the effect of diet type on behavior and present the first time-activity budgets of Lesser White-fronted Geese on natural habitats during the entire wintering period.
引文
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