多空间尺度下驼鹿和狍受人类干扰的生态效应及其适应机制研究
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摘要
自2005年至2007年冬季，综合运用动物生态调查技术、现代景观生态学技术、数学生态学的建模技术、种群生态学、行为生态学等技术以及营养生态学的生化分析技术，在东北小兴安岭西北坡的沾河和胜山地区，布设60条长约2km的样线，调查10m×10m的样方613个，2m×2m的小样方5265个，跟踪足迹链105段53 000m，收集粪便样本184份，检测雪尿样本260份，对多空间尺度下驼鹿(Alces alces cameloides)和狍(Capreolus pygargus bedfordi)受人类干扰的生态效应及其适应机制进行了研究，获得以下6个方面结果：
1、驼鹿和狍在假定无人为干扰和有人为干扰的资源选择模型中，对各种资源变量表现出不同的选择或回避行为。而且，在包括人为干扰变量的模型中，驼鹿和狍对各种人为干扰变量也表现出不同的选择或回避反应。驼鹿在空间分布上不回避狍，模型评估显示两种动物的优质生境都大量丧失，优、良等级生境转化为中、差等级生境，并呈现出相互隔离的斑块，特别是驼鹿受人为活动干扰程度最大。在干扰模型的GIS图层质量分级中，驼鹿的优质生境仅剩一小块孤岛，绝大部分优、良生境转化为差生境，表明一旦人为干扰进一步加重可能导致整个地区驼鹿种群的严重濒危乃至绝迹。
2、驼鹿和狍在研究地区内对可食植物类别的偏爱程度不同。驼鹿对柳，而狍对杨的选择性最高。在不同干扰程度的区域内，驼鹿和狍表现出不同的食物选择性和采食特征；在微生境、斑块和景观尺度下，驼鹿和狍的采食活动呈现出不同的生境需求，而且驼鹿和狍都展示出在微生境尺度对食物丰富度的选择性；人为干扰活动影响驼鹿和狍在各尺度采食活动的资源选择。通过分别建立的斑块和景观尺度模型的GIS图层运算，可知驼鹿和狍优、良等级的生境面积都较中、差等级采食生境的面积小，且平均单位斑块的面积小，生境的丧失及破碎化严重。综合斑块和景观尺度的模型运算，表明整个研究地区内驼鹿采食生境的丧失较狍严重。
3、驼鹿卧息活动在不同尺度条件下呈现出不同的生境资源需求，并且均受公路、居民点和森林采伐时间间隔的影响，且显示出相似的反应距离。由于居民点附近分布的活倒木提供了丰富的食物，吸引驼鹿对其附近区域的选择，对森林采伐时间间隔的延长表现出偏爱性逐渐降低，对公路的回避反应约出现在3.5km；狍的卧息活动也体现出不同尺度下不同的生境需求，并且在这3个尺度均受到居民点的影响，在斑块尺度受到采伐时间间隔的影响。各尺度狍对居民点均显示出相似的反应距离。距居民点6km左右时，狍卧息活动对居民点呈现回避反应。在斑块尺度下，随采伐间隔时间的延长偏爱性增强。
4、应用分维分析获得驼鹿和狍生境利用特征的结果表明，驼鹿移动路径的分维值随着尺度的变化出现明显的间断点，而狍的移动路径随尺度的变化呈直线，表现为自相似性；驼鹿在3个不同分维尺度范围(6-10m，10-15m和6-15m)内的生境利用均与开阔生境的喜好和排粪行为相关，但在6-15m内与寻找食物和对针阔混交林的利用有关，而在6-10m内则与寻找食物和回避针叶林有关；狍在3-20m对生境的利用主要与对密集乔木林的回避以及与加大采食量和采取适宜的卧息策略相关。
5、对驼鹿和狍活动微生境特征的统计和判别分析表明，雪深、当年生柳条比例和采伐时间间隔对判别这两个物种微生境的特征贡献最大。其中，采伐时间间隔对判别驼鹿和狍微生境分离的作用最大，其次是雪深和当年生柳条的比例。驼鹿和狍展示出对森林采伐迹地不同演替阶段特征适应性的差异。因此，适宜的森林采伐干扰活动可能促进两物种微生境的分离，有利共存。
6、驼鹿和狍种群的营养动态研究中，未发现即将死亡的个体(UN:C≥20.0mg:mg)，但发现2006年和2007年1月份沾河地区驼鹿和2006年3月份沾河地区狍种群中都存在较大比例个体受到严重营养限制(UN:C≥3.5mg:mg)，而且2006年和2007年1月至2月胜山地区狍种群的营养状况要好于沾河地区狍。影响沾河地区驼鹿种群营养动态的主要因素可能是与狍之间的食物竞争。当生境较差时，驼鹿和狍均呈现内源蛋白的分解代谢，并通过调整对杨、桦等食物的采食水平来减少与狍的食物重叠，同时采食含有可消化能和粗蛋白更高的冷杉来补充能量和蛋白需求以适应严酷的生境；影响狍种群营养动态的主要因素可能是食物与雪深的共同作用。当生境较差时，狍也靠内源蛋白的分解代谢，同时提高食物中营养成分较高的杨的利用比例及偶而利用草本植物来适应对营养和能量的需求。综合分析食物丰富度、雪深、食性的时空变化与驼鹿和狍种群营养动态之间的关系，发现驼鹿和狍在生境严酷时，各自都呈现出腿部形态(前膝高度)、采食行为、营养代谢等一系列的适应组合。此外，通过细化驼鹿和狍种群个体中UN:C值的分布能够间接反映种群死亡率变化的动态趋势，分析其与生境因子的相关性，使雪尿分析技术在生境日益丧失的鹿类动物种群的保护与管理工作中呈现较高的应用价值。
Ecological effects of anthropogenic disturbances on moose (Alces alces cameloides) and roe deer (Capreolus pygargus bedfordi) at multiple spatial scales were studied by locating 60 2-km transects, surveying 613 10-m×10-m plots and 5265 2-m×2-m subplots, following 105 snow tracks in total length of 53 000-m, collecting 184 fecal samples and assaying 260 snow-urine samples, in the northwestern Lesser Kingan Mountains by using both ecological techniques (such as field survey, modem landscape ecological techniques, mathematic ecological modeling, population ecology techniques, behavioral ecology techniques etc.) and nutritional ecological techniques (such as high performance liquid chromatography and biochemical analysis) from 2005 to 2007. Six conclusions were drawn as follows:
1. In our survey, moose and roe deer exhibited differential selection or avoidance behavior for all kinds of resources and human disturbances. Moose did not appear to avoid the occurrences of roe deer. The spatial distribution of moose revealed no avoidance of roe deer, but differential selection of resources and different responses to human disturbance. Models indicated that the inclusion of human disturbance as a factor in the modelling significantly diminished the assessment of habitat quality, and resulted in an assessment showing an increase in the degree of fragmentation of high quality and good quality habitat patches. Such disturbance was extremely significant for moose, as shown in a comparison of the quality layers of GIS maps with and without the inclusion of human disturbance factors. Most habitat assessed as high quality without the inclusion of any human disturbance factor was assessed as poor quality if human disturbance factors were included in the model; only a small patch of high quality remained. This suggested that human disturbance is a key factor affecting distribution patterns of moose and roe deer.
2. Moose and roe deer exhibited differential preferences for plant species in the whole study area. Moose primarily preferred willows, while roe deer preferred poplars. They exhibited different characteristics of foraging behavior in zones with differing magnitudes of human disturbances. Moose and roe deer appeared to have different habitat requirements for foraging activity at microhabitat, patch and landscape scales, but they all were correlated with food abundances at microhabitat scale. Human disturbance influenced foraging habitat selection of moose and roe deer at the three scales as above. Individual calculations of GIS map layers of patches and landscapes indicated that both the whole area and mean patch area of high and good forage habitats were smaller than that of poor and low ones, suggesting a serious habitat loss and fragmentation. In addition, the loss of moose foraging habitat was more serious than that of roe deer as revealed by combined calculations.
3. For the three different scales, microhabitat, patch and landscape, the moose exhibited different habitat requirements for bed-site selection, but this selection was negatively influenced by proximity to roads, positively influenced by the presence of settlements for forestry workers and researchers, andnegatively influenced by the harvesting interval of the forest patch. It also responded (avoiding or selecting) at similar distances for the three different habitat scales when they were confronted with the same type of human disturbance features. It did not appear to avoid settlement Significantly, in contrast, it showed a slight preference. The avoidance distance for roads occurred at about 3.5km. Habitat preference decreased with the extension of forest harvest intervals. Habitat requirement of roe deer was a little different from that of moose. Bed-site selection of this species was influenced by the proximity of settlements at all three scales, but was only influeced by forest harvest interval at patch scale. The response distance to a settlement was about 6 km at all three scales. In contrast to the moose, habitat preference of roe deer increased with the extension of forest harvest intervals.
4. Fractal analysis of habitat utilization characteristics indicated that fractal value of moose movement path changed with the scales and revealed an apparent abrupt disjoint point, while the fractal value of roe deer showed straight line and self-similarity. These results suggested that moose demonstrated a preference for open habitat and similar fecal pellet excreting behavior at all three fractal scales, but only correlated with foraging in the proximity of food plants and use of mixed coniferous and deciduous forest at 6-15m, and correlated with foraging in the proximity of food plants and avoiding coniferous forest at 6-10m. Roe deer habitat utilization was mainly correlated with avoidance of denser arboreal forest, strategies increasing forage quantity and adopting suitable bedding behavior.
5. Snow depth, the proportion of annual willow shoots and forest harvest intervals were significant variables to differentiate the microhabitat selection of both moose and roe deer. Forest harvest intervals contributed to their microhabitat separation the most, followed by snow depth and the proportion of annual willow shoots. Consequently, both deer species exhibited differences in adaptibility to harvested patches at different successional stages, tolerance to snow depth and preferences in food plant species selection. Hence, suitable forest harvest activities may promote the separation of their microhabitats, benefiting their coexistence.
6. No dead or moribund individuals (UN:C≥20.0mg:mg) were found in these surveys, but a considerable proportion of moose were observed suffering from serious nutritional restriction (UN:C≥3.5mg:mg) in Jan. of 2006 and Jan. 2007 years in Zhanhe region, as well as roe deer in Mar. 2006 in the same region. In contrast, the nutritional condition of roe deer in Shengshan region was better than those in Zhanhe region from Jan. to Feb. in 2006 and 2007. Food competition between moose and roe deer might be a main factor impacting nutritional trends of moose population. At the time that harsh habitat occurred, moose exhibited endogenous protein catabolism. They reduced food overlap with roe deer by regulating foraging frequency for poplar and birch etc. and obtained energy and crude protein by foraging more quantity of faber fir, which has higher energy and protein contents; Snow depth and food abundance might be the main factors impacting the nutritional trends of roe deer population. Roe deer also exhibited endogenous protein catabolism, obtained energy and crude protein met their energy and crude protein requirements by foraging a higher proportion of poplar in their diets (poplar has higher crude protein levels) and a higher proportion of those few grasses which yield more digestible energy, while they were experiencing harsh habitat. Our comprehensive analyses of the relationship between nutritional trend and the temporal and spatial changes of food abundances, snow depths and diets suggested that the two species adopt different series of adaptable combinations of leg morphology, foraging behavior and nutritional metabolism etc. while they were experiencing harsh habitat conditions. In addition, trends of death rate changes of moose and roe deer populations might be indirectly reflected by refining the distribution of UN:C of individuals, which could allow us to analyze its correlation with habitat factors. Therefore, snow-urine analysis might be a valuable tool in practical management of deer populations under conditions of increasing habitat loss.
1、驼鹿和狍在假定无人为干扰和有人为干扰的资源选择模型中，对各种资源变量表现出不同的选择或回避行为。而且，在包括人为干扰变量的模型中，驼鹿和狍对各种人为干扰变量也表现出不同的选择或回避反应。驼鹿在空间分布上不回避狍，模型评估显示两种动物的优质生境都大量丧失，优、良等级生境转化为中、差等级生境，并呈现出相互隔离的斑块，特别是驼鹿受人为活动干扰程度最大。在干扰模型的GIS图层质量分级中，驼鹿的优质生境仅剩一小块孤岛，绝大部分优、良生境转化为差生境，表明一旦人为干扰进一步加重可能导致整个地区驼鹿种群的严重濒危乃至绝迹。
2、驼鹿和狍在研究地区内对可食植物类别的偏爱程度不同。驼鹿对柳，而狍对杨的选择性最高。在不同干扰程度的区域内，驼鹿和狍表现出不同的食物选择性和采食特征；在微生境、斑块和景观尺度下，驼鹿和狍的采食活动呈现出不同的生境需求，而且驼鹿和狍都展示出在微生境尺度对食物丰富度的选择性；人为干扰活动影响驼鹿和狍在各尺度采食活动的资源选择。通过分别建立的斑块和景观尺度模型的GIS图层运算，可知驼鹿和狍优、良等级的生境面积都较中、差等级采食生境的面积小，且平均单位斑块的面积小，生境的丧失及破碎化严重。综合斑块和景观尺度的模型运算，表明整个研究地区内驼鹿采食生境的丧失较狍严重。
3、驼鹿卧息活动在不同尺度条件下呈现出不同的生境资源需求，并且均受公路、居民点和森林采伐时间间隔的影响，且显示出相似的反应距离。由于居民点附近分布的活倒木提供了丰富的食物，吸引驼鹿对其附近区域的选择，对森林采伐时间间隔的延长表现出偏爱性逐渐降低，对公路的回避反应约出现在3.5km；狍的卧息活动也体现出不同尺度下不同的生境需求，并且在这3个尺度均受到居民点的影响，在斑块尺度受到采伐时间间隔的影响。各尺度狍对居民点均显示出相似的反应距离。距居民点6km左右时，狍卧息活动对居民点呈现回避反应。在斑块尺度下，随采伐间隔时间的延长偏爱性增强。
4、应用分维分析获得驼鹿和狍生境利用特征的结果表明，驼鹿移动路径的分维值随着尺度的变化出现明显的间断点，而狍的移动路径随尺度的变化呈直线，表现为自相似性；驼鹿在3个不同分维尺度范围(6-10m，10-15m和6-15m)内的生境利用均与开阔生境的喜好和排粪行为相关，但在6-15m内与寻找食物和对针阔混交林的利用有关，而在6-10m内则与寻找食物和回避针叶林有关；狍在3-20m对生境的利用主要与对密集乔木林的回避以及与加大采食量和采取适宜的卧息策略相关。
5、对驼鹿和狍活动微生境特征的统计和判别分析表明，雪深、当年生柳条比例和采伐时间间隔对判别这两个物种微生境的特征贡献最大。其中，采伐时间间隔对判别驼鹿和狍微生境分离的作用最大，其次是雪深和当年生柳条的比例。驼鹿和狍展示出对森林采伐迹地不同演替阶段特征适应性的差异。因此，适宜的森林采伐干扰活动可能促进两物种微生境的分离，有利共存。
6、驼鹿和狍种群的营养动态研究中，未发现即将死亡的个体(UN:C≥20.0mg:mg)，但发现2006年和2007年1月份沾河地区驼鹿和2006年3月份沾河地区狍种群中都存在较大比例个体受到严重营养限制(UN:C≥3.5mg:mg)，而且2006年和2007年1月至2月胜山地区狍种群的营养状况要好于沾河地区狍。影响沾河地区驼鹿种群营养动态的主要因素可能是与狍之间的食物竞争。当生境较差时，驼鹿和狍均呈现内源蛋白的分解代谢，并通过调整对杨、桦等食物的采食水平来减少与狍的食物重叠，同时采食含有可消化能和粗蛋白更高的冷杉来补充能量和蛋白需求以适应严酷的生境；影响狍种群营养动态的主要因素可能是食物与雪深的共同作用。当生境较差时，狍也靠内源蛋白的分解代谢，同时提高食物中营养成分较高的杨的利用比例及偶而利用草本植物来适应对营养和能量的需求。综合分析食物丰富度、雪深、食性的时空变化与驼鹿和狍种群营养动态之间的关系，发现驼鹿和狍在生境严酷时，各自都呈现出腿部形态(前膝高度)、采食行为、营养代谢等一系列的适应组合。此外，通过细化驼鹿和狍种群个体中UN:C值的分布能够间接反映种群死亡率变化的动态趋势，分析其与生境因子的相关性，使雪尿分析技术在生境日益丧失的鹿类动物种群的保护与管理工作中呈现较高的应用价值。
Ecological effects of anthropogenic disturbances on moose (Alces alces cameloides) and roe deer (Capreolus pygargus bedfordi) at multiple spatial scales were studied by locating 60 2-km transects, surveying 613 10-m×10-m plots and 5265 2-m×2-m subplots, following 105 snow tracks in total length of 53 000-m, collecting 184 fecal samples and assaying 260 snow-urine samples, in the northwestern Lesser Kingan Mountains by using both ecological techniques (such as field survey, modem landscape ecological techniques, mathematic ecological modeling, population ecology techniques, behavioral ecology techniques etc.) and nutritional ecological techniques (such as high performance liquid chromatography and biochemical analysis) from 2005 to 2007. Six conclusions were drawn as follows:
1. In our survey, moose and roe deer exhibited differential selection or avoidance behavior for all kinds of resources and human disturbances. Moose did not appear to avoid the occurrences of roe deer. The spatial distribution of moose revealed no avoidance of roe deer, but differential selection of resources and different responses to human disturbance. Models indicated that the inclusion of human disturbance as a factor in the modelling significantly diminished the assessment of habitat quality, and resulted in an assessment showing an increase in the degree of fragmentation of high quality and good quality habitat patches. Such disturbance was extremely significant for moose, as shown in a comparison of the quality layers of GIS maps with and without the inclusion of human disturbance factors. Most habitat assessed as high quality without the inclusion of any human disturbance factor was assessed as poor quality if human disturbance factors were included in the model; only a small patch of high quality remained. This suggested that human disturbance is a key factor affecting distribution patterns of moose and roe deer.
2. Moose and roe deer exhibited differential preferences for plant species in the whole study area. Moose primarily preferred willows, while roe deer preferred poplars. They exhibited different characteristics of foraging behavior in zones with differing magnitudes of human disturbances. Moose and roe deer appeared to have different habitat requirements for foraging activity at microhabitat, patch and landscape scales, but they all were correlated with food abundances at microhabitat scale. Human disturbance influenced foraging habitat selection of moose and roe deer at the three scales as above. Individual calculations of GIS map layers of patches and landscapes indicated that both the whole area and mean patch area of high and good forage habitats were smaller than that of poor and low ones, suggesting a serious habitat loss and fragmentation. In addition, the loss of moose foraging habitat was more serious than that of roe deer as revealed by combined calculations.
3. For the three different scales, microhabitat, patch and landscape, the moose exhibited different habitat requirements for bed-site selection, but this selection was negatively influenced by proximity to roads, positively influenced by the presence of settlements for forestry workers and researchers, andnegatively influenced by the harvesting interval of the forest patch. It also responded (avoiding or selecting) at similar distances for the three different habitat scales when they were confronted with the same type of human disturbance features. It did not appear to avoid settlement Significantly, in contrast, it showed a slight preference. The avoidance distance for roads occurred at about 3.5km. Habitat preference decreased with the extension of forest harvest intervals. Habitat requirement of roe deer was a little different from that of moose. Bed-site selection of this species was influenced by the proximity of settlements at all three scales, but was only influeced by forest harvest interval at patch scale. The response distance to a settlement was about 6 km at all three scales. In contrast to the moose, habitat preference of roe deer increased with the extension of forest harvest intervals.
4. Fractal analysis of habitat utilization characteristics indicated that fractal value of moose movement path changed with the scales and revealed an apparent abrupt disjoint point, while the fractal value of roe deer showed straight line and self-similarity. These results suggested that moose demonstrated a preference for open habitat and similar fecal pellet excreting behavior at all three fractal scales, but only correlated with foraging in the proximity of food plants and use of mixed coniferous and deciduous forest at 6-15m, and correlated with foraging in the proximity of food plants and avoiding coniferous forest at 6-10m. Roe deer habitat utilization was mainly correlated with avoidance of denser arboreal forest, strategies increasing forage quantity and adopting suitable bedding behavior.
5. Snow depth, the proportion of annual willow shoots and forest harvest intervals were significant variables to differentiate the microhabitat selection of both moose and roe deer. Forest harvest intervals contributed to their microhabitat separation the most, followed by snow depth and the proportion of annual willow shoots. Consequently, both deer species exhibited differences in adaptibility to harvested patches at different successional stages, tolerance to snow depth and preferences in food plant species selection. Hence, suitable forest harvest activities may promote the separation of their microhabitats, benefiting their coexistence.
6. No dead or moribund individuals (UN:C≥20.0mg:mg) were found in these surveys, but a considerable proportion of moose were observed suffering from serious nutritional restriction (UN:C≥3.5mg:mg) in Jan. of 2006 and Jan. 2007 years in Zhanhe region, as well as roe deer in Mar. 2006 in the same region. In contrast, the nutritional condition of roe deer in Shengshan region was better than those in Zhanhe region from Jan. to Feb. in 2006 and 2007. Food competition between moose and roe deer might be a main factor impacting nutritional trends of moose population. At the time that harsh habitat occurred, moose exhibited endogenous protein catabolism. They reduced food overlap with roe deer by regulating foraging frequency for poplar and birch etc. and obtained energy and crude protein by foraging more quantity of faber fir, which has higher energy and protein contents; Snow depth and food abundance might be the main factors impacting the nutritional trends of roe deer population. Roe deer also exhibited endogenous protein catabolism, obtained energy and crude protein met their energy and crude protein requirements by foraging a higher proportion of poplar in their diets (poplar has higher crude protein levels) and a higher proportion of those few grasses which yield more digestible energy, while they were experiencing harsh habitat. Our comprehensive analyses of the relationship between nutritional trend and the temporal and spatial changes of food abundances, snow depths and diets suggested that the two species adopt different series of adaptable combinations of leg morphology, foraging behavior and nutritional metabolism etc. while they were experiencing harsh habitat conditions. In addition, trends of death rate changes of moose and roe deer populations might be indirectly reflected by refining the distribution of UN:C of individuals, which could allow us to analyze its correlation with habitat factors. Therefore, snow-urine analysis might be a valuable tool in practical management of deer populations under conditions of increasing habitat loss.
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