贺兰山岩羊（Pseudois nayaur）和马鹿（Cervus elaphus alxaicus）的食性及生境选择比较研究
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摘要
岩羊(Pseudois nayaur)属偶蹄目、牛科、羊亚科、岩羊属,是青藏高原特有种,在我国重点保护野生动物名录中被列为国家二级保护动物。贺兰山是岩羊的一个重要的集中分布区,20多年来,此区域内岩羊的数量一直呈现出稳定增长的趋势,随着其种群数量的增长,已明显呈现出优势物种的特征,导致同域分布其他有蹄类数量下降甚至濒危,草场退化等问题的出现。贺兰山地区同域分布的有蹄类有岩羊、马鹿(Cervus elaphus alxaicus)和马麝(Moschus chrysogaster),马麝现已相当稀少。由于贺兰山地区是一个“大陆岛屿”,其周围被沙漠、城市和河流(黄河)所阻断,贺兰山的有蹄类动物种群实际上是隔离种群,种群数量的波动不受迁入和迁出的影响,而种间竞争是影响其生存的一个重要因素。因此,本研究以贺兰山同域分布的岩羊和马鹿为对象,对比分析了岩羊和马鹿在食性、生境选择方面的差异,并在此基础上对二者共存机制进行初步探讨,这对贺兰山生态系统的稳定和生物多样性的保护具有重要的探索意义,也为野生动物及其栖息地的保护和管理工作提供科学依据和理论指导。
2006年至2008年,在贺兰山地区对同域分布岩羊和马鹿活动区域里两种动物采食的植物种类进行了调查,并采集了岩羊和马鹿活动区域内可采食的植物标本以及岩羊和马鹿粪便,利用粪便显微分析法对二者各个季节的食性和二者在食性方面的差异性进行了研究分析；同时,利用痕迹检测法和直接观察法对同域分布的岩羊和马鹿各个季节的生境选择进行了调查研究,分别分析了它们各自在不同季节选择和利用取食生境和卧息生境的差异性,并深入对比分析了二者在生境选择方面的差异。每个季节从15个沟系采集岩羊和马鹿粪便复合样本各60个,共组成480个样本;同时测定了岩羊利用样方804个,其中取食利用样方518个,卧息利用样方286个；测定马鹿利用样方661个,其中取食利用样方341个,卧息利用样方320个。对岩羊利用生境样方设定对照样方共202个,对马鹿利用生境样方设定对照样方共707个。主要研究结果如下：
(1)对贺兰山地区岩羊的食物组成、季节性变化进行了研究。结果表明,贺兰山岩羊全年共取食植物类别共有25科54种(属)植物。采食各科植物中禾本科(49.48%)、榆科(18.15%)、蔷薇科(16.61%)所占比例较高,组成了岩羊全年的主要食物。其中针茅、灰榆、早熟禾在4个季节里均具有较高的RD值,总量为全年取食植物组成的60.24%,为岩羊全年的大宗食物。岩羊在取食植物种类上存在季节性差异,春、夏、秋季均大量采食禾本科草本植物,全年平均取食量为49.48%；冬季以灌木和乔木为主,取食非禾本科草本植物全年平均在10.78%。在取食多样性方面,夏季取食植物种类35种(属),Shannon-Wiener指数H'冬季高于其他3个季节；食物生态位宽度B值冬季和夏季高于其他2个季节,并且冬季B值略高于夏季。
(2)对贺兰山地区马鹿的食物组成、季节性变化进行了研究。贺兰山马鹿全年取食植物种类为25科58种(属)。杨柳科(36.24%)、榆科(19.72%)、蔷薇科(13.52%)与禾本科(13.49%)植物为马鹿4个季节的主要食物。其中,灰榆、小红柳、山杨、早熟禾和针茅4个季节的RD平均值分别为19.72%、19.53%、16.32%、6.29%和6.05%,占马鹿全年食物组成比例的67.91%。贺兰山马鹿在取食植物类别上也存在一定差异,全年均大量取食乔木类植物,RD平均达到60.43%,其次是灌木和禾本科草本植物,较少取食非禾本科草本植物。贺兰山马鹿各个季节取食多样性、均匀度指数和生态位宽度也存在差异,夏季取食食物种类最多,为44种；其次为春季和秋季,冬季仅为23种,为4个季节最少；均匀度指数在冬季最高为1.198,其次为秋季,夏季最低为0.700。冬季的食物生态位宽度远高于其他3个季节,为8.234,春季最低,为4.097。
(3)对贺兰山同域分布的岩羊和马鹿食性进行了对比分析。结果表明,贺兰山同域分布的岩羊和马鹿食性存在较大差异。具体表现在全年主要取食食物上,虽然针茅、灰榆和早熟禾在岩羊和马鹿全年平均取食量中均有较高比例,但是取食的季节性差异较大。在全年取食植物的科别上,岩羊主要取食禾本科、榆科、蔷薇科和伞形科植物,而马鹿主要以杨柳科、榆科、蔷薇科、禾本科和豆科为主。二者对杨柳科和禾本科的取食差别很大,岩羊很少取食杨柳科(1.01%),大量取食禾本科(49.48%),而马鹿取食杨柳科的比例达到36.24%,禾本科为13.49%。另外,岩羊和马鹿对不同类别植物的取食也有一定的差异,岩羊取食较高比例的禾本科草本(49.48%),其次是乔木和灌木；而马鹿大量取食乔木,比例高达60.43%,其次为灌木和禾本科草本,二者对非禾本科草本均少量取食。同时,岩羊和马鹿在各季节均有大量食物重叠,但较高的重叠指数只出现在冬春两季,并且大宗食物中重叠数量很少。
(4)利用直接观察法和痕迹检测法对贺兰山岩羊4个季节的取食和卧息生境的进行了测定,并利用卡方检验、Kolmogorov-Smirnov检验、kruskal-Wallis H检验、主成分分析等方法对不同季节岩羊取食和卧息生境选择和利用及其差异性进行了研究。研究发现,岩羊四季均偏爱选择山地疏林草原带,海拔在2000 m以下有明显断裂的山坡上,接近水源和裸岩,对取食和卧息生境中坡向和坡度几乎无选择性。岩羊取食活动更多集中在灰榆林中,偏爱取食灰榆嫩枝和树叶,生境内乔木稀疏矮小,且距离较远,灌木密度较大、且高度在1.3 m以上,食物丰盛度高。贺兰山岩羊卧息生境存在较大的季节性差异,春夏秋季多选择在山地疏林草原带,冬季由于高海拔地区积雪原因,多到低海拔地区的山地草原带活动,卧息地乔木稀疏,高度一般在4 m以下,距离较远,对灌木多随机选择或无选择,生境远离人为干扰。
(5)利用上述方法对贺兰山马鹿四季取食和卧息生境进行了测定,并对其不同季节生境选择和利用的差异性进行了研究。研究发现,贺兰山马鹿多活动于海拔高度在2000 m以上平滑起伏或中等断裂的山坡,活动存在明显的季节性迁徙现象,因此在取食和卧息生境选择上具有显著的季节性差异。山地草原带和山地针叶林带是马鹿阿拉善亚种在贺兰山所选择的主要植被类型,随季节变化而迁徙,冬春季多活动于低海拔地区的山地草原带,选择向阳的山坡；夏秋季偏爱选择高海拔地区的山地针叶林地带,多选择半阴半阳或阴坡活动；偏爱具有高大、浓密灌木,远离裸岩的生境,并且对隐蔽级要求较高。坡度是贺兰山马鹿生境选择的一个重要限制因子,在整个研究区域贺兰山马鹿通常选择小于10°的缓坡。而对距水源距离和人为干扰距离几乎无选择性。
(6)利用卡方检验、Mann-Whitey U检验及逐步判别分析等方法对贺兰山岩羊和马鹿各季节的生境选择差异性进行了对比研究。在所设定的17种生态因子中,典则判别系数和逐步判别分析的结果显示贺兰山同域分布的岩羊和马鹿各个季节在取食和卧息生境的选择上几乎完全分离,也表明不同季节岩羊和马鹿对取食和卧息生境的选择存在显著的差异。卡方检验结果表明贺兰山岩羊和马鹿不同季节在取食和卧息生境选择上对植被类型、地形特征、优势乔木和坡向等4种非数值型生态因子的利用均存在极显著差异(P<0.01),Mann-Whitey U检验表明,贺兰山岩羊与马鹿4个季节的取食生境在乔木密度、乔木高度、灌木密度、灌木高度、食物丰盛度、坡度、隐蔽级、人为干扰距离和距裸岩距离等生态因子上具有极显著差异(P<0.01)或显著差异(P<0.05),而卧息生境在灌木密度、食物丰盛度、坡度、隐蔽级、距裸岩距离、海拔高度和人为干扰距离等生态因子上具有极显著(P<0.01)或显著的差异(P<0.05)。从而也说明贺兰山岩羊和马鹿在各个季节有明显的生境分离,尤其是在食物资源比较匮乏的条件下,通过生境分离来实现资源的平衡分配是岩羊和马鹿的选择策略之一,这与预测吻合。
(7)对贺兰山同域分布的岩羊和马鹿的竞争—共存机制进行了初步探讨。结果表明贺兰山地区同域分布的岩羊和马鹿对食物资源和生境资源的利用存在一定的竞争,二者通过以下途径实现共存：①岩羊和马鹿通过资源生态位的分化来实现共存,二者在食性和采食对策上存在较大的分离,包括对食物种类和取食地的选择两个途径：②岩羊和马鹿通过不同的生境利用方式来适应环境、降低竞争,从而达到同域共存。
Blue sheep (Pseudois nayaur) is one of Qinghai-Tibet Plateau endemic ungulates belong to the sunfamily Caprinae gunus Pseudois (Artiodactyla, Bovidae). It was listed in the II grade nationally protected animals of China. Helan Mountain is an important distribution area of blue sheep; their populations increased steadily in the past 20 years, and becoming a dominant species here. The increase of blue sheep's population resulted in a decrease of population in other ungulates and grassland degradation. Ungulates sympatric in Helan Mountain are blue sheep, red deer (Cervus elaphus alxaicus), and moschus sifanicus(Moschus chrysogaster). The population of ungulates in this region is believed to be an isolated geographical population, because the Helan Mountain is surrounded by Yinchuan City, deserts and Yellow river, and the competition among sympatric ungulates affected their survive. In this study, we analyzed the differences of diet and habitat selection by blue sheep and red deer, and made a further study on the coexistence mechanisms of them.
From 2006 to 2008, we collected and studied the vegetation species consumed by sympatric blue sheep(Pseudois nayaur) and red deer(Cervus elaphus alxaicus) and their fecal samples in the Helan Mountains. The annual food component and proportion of blue sheep and red deer were studied with the fecal micro histological analysis. The habitat selection of sympatric blue sheep and red deer, especially the difference of them were studied based on direct observation and examination of fresh sites used by blue sheep and red deer throughout the study area. A total number of 480 composite samples of fresh feces of blue sheep and red deer were collected from 15 valleys.804 sites composed of 518 feeding and 286 bedding used by blue sheep and 661 sites composed of 341 feeding and 320 bedding used by deer were measured in the study area,202 random sites for the sites used by blue sheep and 707 random sites for the sites used by red deer were also measured, and the main results are as follows:
(1) The food composition and diet seasonal changes of blue sheep were studied, results showed that blue sheep(Pseudios nayaur) in Helan mountains consumed 54 plant species (genera) belonging to 25 families in the whole year. In the diet of composition, the main families blue sheep foraged were Gramineae, Rosaceae, Ulmaceae and Leguminosae, making up the main food in the whole year. And Stipa spp. Poa spp.and Ulmus glaucescens that had high RD in the diet composition were most common and dominated the diet of blue sheep. Plant species blue sheep foraged revealed seasonal difference.35 plant species (genera) were identified in the microhistological analyses in summer, more than those in other three seasons. Shannon-Wiener H'was the highest in the four seasons. Index of diet niche in winter was the highest, followed by that in summer.
(2) The food composition and diet seasonal changes of red deer were also studied, and results showed plant species red deer(Cerus elaphus alxaicus) in Helan Mountains foraged were 58 species (genera) from 25 families. We sonsidered red deer as polyphagous herbivores. In the whole year, Salicaceae, Rosaceae, Salicaceae and Gramineae in the diet composition made up 83.53%, were main food. The average RD of Salicaceae in the diet was 33.44%, followed by Rosaceae (19.99%), Salicaceae (15.22%) and Gramineae (15.13%). The average Rd of Salix microtachya var. bordensis, Ulmus glaucescens, Populus davidiana and Stipa spp. were 19.92%、16.11%、13.80%and 8.55%separately, making up 58.38%of diet composition of red deer. The characteristic of different plant category red deer showed seasonal variance. The sequence of plant category was trees (RD; 55.51%), shrubs (RD; 22.47%), grminoids (RD; 17.32%) and forbs (RD; 4.71%). The highest proportion of tree during the microhistological analyses on red deer fecal was in summer, making up the 89.27%of the diet composition. Shrubs made up 44.70%of the diet composition in winter. RD of grminoids in autumn was 56.38%and forbs 8.0%, lowest. Results showed that plant species, Pielou index, and index of diet niche were different among the four seasons. Plant species red deer foraged were 44, followed by spring and autumn, winter (22). Shannon-Wiener index of winter was the highest than that in the other three seasons. Pielou index in winter also was the highest,1.198, followed by autumn, and index of summer was the lowest. B, index of diet niche, was 8.234 in winter and 4.097 in spring.
(3) Blue sheep and red deer in Helan mountains were sympatric, but the diet composition of them were significantly different, especially the main food of the two ungulates. Even though the Ulmus glaucescens, Stipa spp. and Poa spp. made up high proportion all the year in the diet of blue sheep and red deer, the RD of them revealed significant seasonal difference. Blue sheep foraged much gramineae, ulmaceae, rosaceae and umbelliferae. But red deer used more Salicaceae, ulmaceae, rosaceae, gramineae and leguminosae. The two ungulates had difference when they foraged salicaceae and gramineae. Blue sheep ate less salicaceae (1.01%) and more gramineae (49.48%). While red deer foraged more salicaceae (36.24%) and less gramineae (13.49%). The proportion of different plant category was different. Blue sheep used high proportion of graminoid (49.48%), followed by tree and shrub. Red deer foraged more tree (60.43%), followed by shrub and graminoid. Forbs were the least. And much food were overlap between blue sheep and red deer. High overlap index emerged in winter and spring, and main food was not overlap significantly.
(4) The feeding and bedding habitat used by blue sheep were measured with the methods of fresh trace observation, assisting by direct observation in 4 seasons, and the data were analyzed by Chi-Square test, Kolmogorov-Smirnov Test, Bonferroni, Kruskal-Wallis H Test and PCA. The result showed that blue sheep preferred to use montane savanna dominated by Ulmus glaucescens when they feeding and bedding, and showed no significant preference to slope and direction, but activity mostly centered in higher slope habitat. Blue sheep preferred to select habitat close to bare rock in summer, autumn and winter, but not in spring. Blue sheep preferred to use habitats dominated by Ulmus glaucescens and dense high shrub as feeding, mostly in distinctly broken slope, close to bare rocks. They used Ulmus glaucescens under 4 m more frequently in summer and autumn, but used mixed trees include Ulmus glaucescens and some conifer below 6 m more in winter and spring. There were distinct differences in the bedding habitat selection of blue sheep in different seasons, they preferred to use montane savanna in spring, summer and autumn, but moved to montane grassland due to heavy snow in higher regions. Their bedding habitat required fewer shrub and fewer trees below 4 m, far from the bedding place.
(5) The habitat selected by red deer (Cervus elaphus alxaicus) was analyzed with the same methods as blue sheep habitat selection. The result showed that red deer preferred to use habitat in gently smooth undulating slopes at higher altitude of more than 2000 m, and they migrated as season changing. There were significant seasonal changes in red deer's habitat selection for feeding and bedding. Red deer preferred montane grassland distributed in lower altitude in spring and winter, but montane conifer forest at higher altitude in summer and autumn. They preferred habitat with high and dense shrub, far from bare rock and high coverage. The slope was an important restrict factor for red deer in Helan Mountain regions, they preferred to use habitats in gentle slope within 10°.But they showed non-selective to the factors of distance to water resource and human disturbance.
(6) The difference in habitat selection of blue sheep and red deer in different seasons was analyzed with chi-square test, Mann-Whitey U test and stepwise discriminant analysis. The result of stepwise discriminant and canonical discriminant coefficients showed that 17 eco-factors in feeding and bedding habitats used by blue sheep and red deer were almost completely separated, indicating a significant difference in their habitat selections. The result of chi-square test described an extremely significant difference (P < 0.01) between blue sheep and red deer in the using of vegetation types, topography, dominant tree and slope aspect. Mann-Whitey U test showed that blue sheep and red deer used tree density, tree height, shrub density, shrub height, food abundance, slope, hiding cover, distance to human disturbance and the distance to bare rock differently at extremely significant level (P<0.01) or significant level (P<0.05) in feeding habitat; and used shrub density, food abundance, slope, hiding cover, distance to bare rock, altitude and distance to human disturbance differently at extremely significant level (P<0.01) or significant level (P<0.05) in bedding habitat. The results showed that there were significant differences in habitat selections between blue sheep and red deer among 4 seasons, especially when the food resources was deficient. The habitat segregation is one of their strategies to meet an equal sharing of resources, which was consistent with our prediction.
(7) The competition-coexistence mechanisms of sympatric distributed blue sheep and red deer in Helan Mountains was discussed. The result showed competition between blue sheep and red deer sympatric distributed in Helan Mountains in the using of food and space resources. They got co-existence with followed means:①Blue sheep and red deer co-existed by means of resource niche differentiation, there were great separations in diet and foraging strategies, including food types, foraging time and foraging habitat selection.②Blue sheep and red deer adapted to the environment and reduced competition in using different habitat patterns, to achieve benign symbiotic relationship in the same area.
2006年至2008年,在贺兰山地区对同域分布岩羊和马鹿活动区域里两种动物采食的植物种类进行了调查,并采集了岩羊和马鹿活动区域内可采食的植物标本以及岩羊和马鹿粪便,利用粪便显微分析法对二者各个季节的食性和二者在食性方面的差异性进行了研究分析；同时,利用痕迹检测法和直接观察法对同域分布的岩羊和马鹿各个季节的生境选择进行了调查研究,分别分析了它们各自在不同季节选择和利用取食生境和卧息生境的差异性,并深入对比分析了二者在生境选择方面的差异。每个季节从15个沟系采集岩羊和马鹿粪便复合样本各60个,共组成480个样本;同时测定了岩羊利用样方804个,其中取食利用样方518个,卧息利用样方286个；测定马鹿利用样方661个,其中取食利用样方341个,卧息利用样方320个。对岩羊利用生境样方设定对照样方共202个,对马鹿利用生境样方设定对照样方共707个。主要研究结果如下：
(1)对贺兰山地区岩羊的食物组成、季节性变化进行了研究。结果表明,贺兰山岩羊全年共取食植物类别共有25科54种(属)植物。采食各科植物中禾本科(49.48%)、榆科(18.15%)、蔷薇科(16.61%)所占比例较高,组成了岩羊全年的主要食物。其中针茅、灰榆、早熟禾在4个季节里均具有较高的RD值,总量为全年取食植物组成的60.24%,为岩羊全年的大宗食物。岩羊在取食植物种类上存在季节性差异,春、夏、秋季均大量采食禾本科草本植物,全年平均取食量为49.48%；冬季以灌木和乔木为主,取食非禾本科草本植物全年平均在10.78%。在取食多样性方面,夏季取食植物种类35种(属),Shannon-Wiener指数H'冬季高于其他3个季节；食物生态位宽度B值冬季和夏季高于其他2个季节,并且冬季B值略高于夏季。
(2)对贺兰山地区马鹿的食物组成、季节性变化进行了研究。贺兰山马鹿全年取食植物种类为25科58种(属)。杨柳科(36.24%)、榆科(19.72%)、蔷薇科(13.52%)与禾本科(13.49%)植物为马鹿4个季节的主要食物。其中,灰榆、小红柳、山杨、早熟禾和针茅4个季节的RD平均值分别为19.72%、19.53%、16.32%、6.29%和6.05%,占马鹿全年食物组成比例的67.91%。贺兰山马鹿在取食植物类别上也存在一定差异,全年均大量取食乔木类植物,RD平均达到60.43%,其次是灌木和禾本科草本植物,较少取食非禾本科草本植物。贺兰山马鹿各个季节取食多样性、均匀度指数和生态位宽度也存在差异,夏季取食食物种类最多,为44种；其次为春季和秋季,冬季仅为23种,为4个季节最少；均匀度指数在冬季最高为1.198,其次为秋季,夏季最低为0.700。冬季的食物生态位宽度远高于其他3个季节,为8.234,春季最低,为4.097。
(3)对贺兰山同域分布的岩羊和马鹿食性进行了对比分析。结果表明,贺兰山同域分布的岩羊和马鹿食性存在较大差异。具体表现在全年主要取食食物上,虽然针茅、灰榆和早熟禾在岩羊和马鹿全年平均取食量中均有较高比例,但是取食的季节性差异较大。在全年取食植物的科别上,岩羊主要取食禾本科、榆科、蔷薇科和伞形科植物,而马鹿主要以杨柳科、榆科、蔷薇科、禾本科和豆科为主。二者对杨柳科和禾本科的取食差别很大,岩羊很少取食杨柳科(1.01%),大量取食禾本科(49.48%),而马鹿取食杨柳科的比例达到36.24%,禾本科为13.49%。另外,岩羊和马鹿对不同类别植物的取食也有一定的差异,岩羊取食较高比例的禾本科草本(49.48%),其次是乔木和灌木；而马鹿大量取食乔木,比例高达60.43%,其次为灌木和禾本科草本,二者对非禾本科草本均少量取食。同时,岩羊和马鹿在各季节均有大量食物重叠,但较高的重叠指数只出现在冬春两季,并且大宗食物中重叠数量很少。
(4)利用直接观察法和痕迹检测法对贺兰山岩羊4个季节的取食和卧息生境的进行了测定,并利用卡方检验、Kolmogorov-Smirnov检验、kruskal-Wallis H检验、主成分分析等方法对不同季节岩羊取食和卧息生境选择和利用及其差异性进行了研究。研究发现,岩羊四季均偏爱选择山地疏林草原带,海拔在2000 m以下有明显断裂的山坡上,接近水源和裸岩,对取食和卧息生境中坡向和坡度几乎无选择性。岩羊取食活动更多集中在灰榆林中,偏爱取食灰榆嫩枝和树叶,生境内乔木稀疏矮小,且距离较远,灌木密度较大、且高度在1.3 m以上,食物丰盛度高。贺兰山岩羊卧息生境存在较大的季节性差异,春夏秋季多选择在山地疏林草原带,冬季由于高海拔地区积雪原因,多到低海拔地区的山地草原带活动,卧息地乔木稀疏,高度一般在4 m以下,距离较远,对灌木多随机选择或无选择,生境远离人为干扰。
(5)利用上述方法对贺兰山马鹿四季取食和卧息生境进行了测定,并对其不同季节生境选择和利用的差异性进行了研究。研究发现,贺兰山马鹿多活动于海拔高度在2000 m以上平滑起伏或中等断裂的山坡,活动存在明显的季节性迁徙现象,因此在取食和卧息生境选择上具有显著的季节性差异。山地草原带和山地针叶林带是马鹿阿拉善亚种在贺兰山所选择的主要植被类型,随季节变化而迁徙,冬春季多活动于低海拔地区的山地草原带,选择向阳的山坡；夏秋季偏爱选择高海拔地区的山地针叶林地带,多选择半阴半阳或阴坡活动；偏爱具有高大、浓密灌木,远离裸岩的生境,并且对隐蔽级要求较高。坡度是贺兰山马鹿生境选择的一个重要限制因子,在整个研究区域贺兰山马鹿通常选择小于10°的缓坡。而对距水源距离和人为干扰距离几乎无选择性。
(6)利用卡方检验、Mann-Whitey U检验及逐步判别分析等方法对贺兰山岩羊和马鹿各季节的生境选择差异性进行了对比研究。在所设定的17种生态因子中,典则判别系数和逐步判别分析的结果显示贺兰山同域分布的岩羊和马鹿各个季节在取食和卧息生境的选择上几乎完全分离,也表明不同季节岩羊和马鹿对取食和卧息生境的选择存在显著的差异。卡方检验结果表明贺兰山岩羊和马鹿不同季节在取食和卧息生境选择上对植被类型、地形特征、优势乔木和坡向等4种非数值型生态因子的利用均存在极显著差异(P<0.01),Mann-Whitey U检验表明,贺兰山岩羊与马鹿4个季节的取食生境在乔木密度、乔木高度、灌木密度、灌木高度、食物丰盛度、坡度、隐蔽级、人为干扰距离和距裸岩距离等生态因子上具有极显著差异(P<0.01)或显著差异(P<0.05),而卧息生境在灌木密度、食物丰盛度、坡度、隐蔽级、距裸岩距离、海拔高度和人为干扰距离等生态因子上具有极显著(P<0.01)或显著的差异(P<0.05)。从而也说明贺兰山岩羊和马鹿在各个季节有明显的生境分离,尤其是在食物资源比较匮乏的条件下,通过生境分离来实现资源的平衡分配是岩羊和马鹿的选择策略之一,这与预测吻合。
(7)对贺兰山同域分布的岩羊和马鹿的竞争—共存机制进行了初步探讨。结果表明贺兰山地区同域分布的岩羊和马鹿对食物资源和生境资源的利用存在一定的竞争,二者通过以下途径实现共存：①岩羊和马鹿通过资源生态位的分化来实现共存,二者在食性和采食对策上存在较大的分离,包括对食物种类和取食地的选择两个途径：②岩羊和马鹿通过不同的生境利用方式来适应环境、降低竞争,从而达到同域共存。
Blue sheep (Pseudois nayaur) is one of Qinghai-Tibet Plateau endemic ungulates belong to the sunfamily Caprinae gunus Pseudois (Artiodactyla, Bovidae). It was listed in the II grade nationally protected animals of China. Helan Mountain is an important distribution area of blue sheep; their populations increased steadily in the past 20 years, and becoming a dominant species here. The increase of blue sheep's population resulted in a decrease of population in other ungulates and grassland degradation. Ungulates sympatric in Helan Mountain are blue sheep, red deer (Cervus elaphus alxaicus), and moschus sifanicus(Moschus chrysogaster). The population of ungulates in this region is believed to be an isolated geographical population, because the Helan Mountain is surrounded by Yinchuan City, deserts and Yellow river, and the competition among sympatric ungulates affected their survive. In this study, we analyzed the differences of diet and habitat selection by blue sheep and red deer, and made a further study on the coexistence mechanisms of them.
From 2006 to 2008, we collected and studied the vegetation species consumed by sympatric blue sheep(Pseudois nayaur) and red deer(Cervus elaphus alxaicus) and their fecal samples in the Helan Mountains. The annual food component and proportion of blue sheep and red deer were studied with the fecal micro histological analysis. The habitat selection of sympatric blue sheep and red deer, especially the difference of them were studied based on direct observation and examination of fresh sites used by blue sheep and red deer throughout the study area. A total number of 480 composite samples of fresh feces of blue sheep and red deer were collected from 15 valleys.804 sites composed of 518 feeding and 286 bedding used by blue sheep and 661 sites composed of 341 feeding and 320 bedding used by deer were measured in the study area,202 random sites for the sites used by blue sheep and 707 random sites for the sites used by red deer were also measured, and the main results are as follows:
(1) The food composition and diet seasonal changes of blue sheep were studied, results showed that blue sheep(Pseudios nayaur) in Helan mountains consumed 54 plant species (genera) belonging to 25 families in the whole year. In the diet of composition, the main families blue sheep foraged were Gramineae, Rosaceae, Ulmaceae and Leguminosae, making up the main food in the whole year. And Stipa spp. Poa spp.and Ulmus glaucescens that had high RD in the diet composition were most common and dominated the diet of blue sheep. Plant species blue sheep foraged revealed seasonal difference.35 plant species (genera) were identified in the microhistological analyses in summer, more than those in other three seasons. Shannon-Wiener H'was the highest in the four seasons. Index of diet niche in winter was the highest, followed by that in summer.
(2) The food composition and diet seasonal changes of red deer were also studied, and results showed plant species red deer(Cerus elaphus alxaicus) in Helan Mountains foraged were 58 species (genera) from 25 families. We sonsidered red deer as polyphagous herbivores. In the whole year, Salicaceae, Rosaceae, Salicaceae and Gramineae in the diet composition made up 83.53%, were main food. The average RD of Salicaceae in the diet was 33.44%, followed by Rosaceae (19.99%), Salicaceae (15.22%) and Gramineae (15.13%). The average Rd of Salix microtachya var. bordensis, Ulmus glaucescens, Populus davidiana and Stipa spp. were 19.92%、16.11%、13.80%and 8.55%separately, making up 58.38%of diet composition of red deer. The characteristic of different plant category red deer showed seasonal variance. The sequence of plant category was trees (RD; 55.51%), shrubs (RD; 22.47%), grminoids (RD; 17.32%) and forbs (RD; 4.71%). The highest proportion of tree during the microhistological analyses on red deer fecal was in summer, making up the 89.27%of the diet composition. Shrubs made up 44.70%of the diet composition in winter. RD of grminoids in autumn was 56.38%and forbs 8.0%, lowest. Results showed that plant species, Pielou index, and index of diet niche were different among the four seasons. Plant species red deer foraged were 44, followed by spring and autumn, winter (22). Shannon-Wiener index of winter was the highest than that in the other three seasons. Pielou index in winter also was the highest,1.198, followed by autumn, and index of summer was the lowest. B, index of diet niche, was 8.234 in winter and 4.097 in spring.
(3) Blue sheep and red deer in Helan mountains were sympatric, but the diet composition of them were significantly different, especially the main food of the two ungulates. Even though the Ulmus glaucescens, Stipa spp. and Poa spp. made up high proportion all the year in the diet of blue sheep and red deer, the RD of them revealed significant seasonal difference. Blue sheep foraged much gramineae, ulmaceae, rosaceae and umbelliferae. But red deer used more Salicaceae, ulmaceae, rosaceae, gramineae and leguminosae. The two ungulates had difference when they foraged salicaceae and gramineae. Blue sheep ate less salicaceae (1.01%) and more gramineae (49.48%). While red deer foraged more salicaceae (36.24%) and less gramineae (13.49%). The proportion of different plant category was different. Blue sheep used high proportion of graminoid (49.48%), followed by tree and shrub. Red deer foraged more tree (60.43%), followed by shrub and graminoid. Forbs were the least. And much food were overlap between blue sheep and red deer. High overlap index emerged in winter and spring, and main food was not overlap significantly.
(4) The feeding and bedding habitat used by blue sheep were measured with the methods of fresh trace observation, assisting by direct observation in 4 seasons, and the data were analyzed by Chi-Square test, Kolmogorov-Smirnov Test, Bonferroni, Kruskal-Wallis H Test and PCA. The result showed that blue sheep preferred to use montane savanna dominated by Ulmus glaucescens when they feeding and bedding, and showed no significant preference to slope and direction, but activity mostly centered in higher slope habitat. Blue sheep preferred to select habitat close to bare rock in summer, autumn and winter, but not in spring. Blue sheep preferred to use habitats dominated by Ulmus glaucescens and dense high shrub as feeding, mostly in distinctly broken slope, close to bare rocks. They used Ulmus glaucescens under 4 m more frequently in summer and autumn, but used mixed trees include Ulmus glaucescens and some conifer below 6 m more in winter and spring. There were distinct differences in the bedding habitat selection of blue sheep in different seasons, they preferred to use montane savanna in spring, summer and autumn, but moved to montane grassland due to heavy snow in higher regions. Their bedding habitat required fewer shrub and fewer trees below 4 m, far from the bedding place.
(5) The habitat selected by red deer (Cervus elaphus alxaicus) was analyzed with the same methods as blue sheep habitat selection. The result showed that red deer preferred to use habitat in gently smooth undulating slopes at higher altitude of more than 2000 m, and they migrated as season changing. There were significant seasonal changes in red deer's habitat selection for feeding and bedding. Red deer preferred montane grassland distributed in lower altitude in spring and winter, but montane conifer forest at higher altitude in summer and autumn. They preferred habitat with high and dense shrub, far from bare rock and high coverage. The slope was an important restrict factor for red deer in Helan Mountain regions, they preferred to use habitats in gentle slope within 10°.But they showed non-selective to the factors of distance to water resource and human disturbance.
(6) The difference in habitat selection of blue sheep and red deer in different seasons was analyzed with chi-square test, Mann-Whitey U test and stepwise discriminant analysis. The result of stepwise discriminant and canonical discriminant coefficients showed that 17 eco-factors in feeding and bedding habitats used by blue sheep and red deer were almost completely separated, indicating a significant difference in their habitat selections. The result of chi-square test described an extremely significant difference (P < 0.01) between blue sheep and red deer in the using of vegetation types, topography, dominant tree and slope aspect. Mann-Whitey U test showed that blue sheep and red deer used tree density, tree height, shrub density, shrub height, food abundance, slope, hiding cover, distance to human disturbance and the distance to bare rock differently at extremely significant level (P<0.01) or significant level (P<0.05) in feeding habitat; and used shrub density, food abundance, slope, hiding cover, distance to bare rock, altitude and distance to human disturbance differently at extremely significant level (P<0.01) or significant level (P<0.05) in bedding habitat. The results showed that there were significant differences in habitat selections between blue sheep and red deer among 4 seasons, especially when the food resources was deficient. The habitat segregation is one of their strategies to meet an equal sharing of resources, which was consistent with our prediction.
(7) The competition-coexistence mechanisms of sympatric distributed blue sheep and red deer in Helan Mountains was discussed. The result showed competition between blue sheep and red deer sympatric distributed in Helan Mountains in the using of food and space resources. They got co-existence with followed means:①Blue sheep and red deer co-existed by means of resource niche differentiation, there were great separations in diet and foraging strategies, including food types, foraging time and foraging habitat selection.②Blue sheep and red deer adapted to the environment and reduced competition in using different habitat patterns, to achieve benign symbiotic relationship in the same area.
引文
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