陆生食肉动物竞争与共存研究概述

陆生食肉动物(食肉目哺乳动物, 以下简称食肉动物)作为食物链与营养级的高位物种对维持生态系统结构与功能稳定性起到重要作用。过度人类干扰已在全球范围内造成食肉动物种群数量剧烈下降和栖息地质量显著退化, 探究食肉动物的区域共存机制对理解生物群落构建、濒危物种保护与管理具有重要意义。本文通过梳理100余篇有关食肉动物在空间、时间和营养3个生态位维度上相互作用的研究, 分析了体型大小、猎物组成、种群结构、环境差异、人类干扰和气候变化等因素对食肉动物种间关系和区域共存的影响, 并对今后食肉动物区域共存研究中亟需解决的问题进行了展望。食肉动物通过生态位分离达到共存并没有单一的理论解释, 猎物、栖息地和人类干扰等因素可以调节食肉动物相互作用关系并直接或间接地影响共存, 共同适应在食肉动物区域共存中具重要作用。食肉动物区域共存是经过长期演化形成的相对稳定状态, 需要以动态的眼光去审视。要明晰生态位重叠与区域共存机制的区别与联系, 在理解生态位分离的基础上, 结合生活史、家域和行为等对食肉动物共存进行综合分析。     

四川卧龙国家级自然保护区三种高山同域鸡形目鸟类的时空生态位比较

高海拔山区气候条件恶劣, 资源匮乏, 探究同域分布的近缘物种如何利用有限的资源以实现稳定共存, 对于了解高山生态系统生物多样性格局的形成和维持机制具有重要意义。鸡形目鸟类飞行能力弱, 属于典型的地栖物种, 生态位空间相对狭窄, 可能面临更高的种间竞争压力。本研究旨在比较几种同域分布的鸡形目鸟类的时空生态位, 为了解高山生态系统同域物种的共存机制提供新的研究案例。2020年4-9月, 研究人员在四川卧龙国家级自然保护区海拔3,300-4,200 m的高山区域进行了野外调查, 通过样线法和样方法对鸡形目鸟类群落优势物种绿尾虹雉(Lophophorus lhuysii)、雉鹑(Tetraophasis obscurus)和雪鹑(Lerwa lerwa)繁殖期的微生境进行调查, 使用红外相机对其活动节律进行监测, 并运用核密度估计法从微生境利用和日活动节律两个生态维度进行了种间生态位比较。结果显示, 雪鹑在微生境利用和日活动节律上均与其他两个物种存在显著差异。绿尾虹雉与雉鹑在微生境的利用上具有相似偏好; 但绿尾虹雉的早活动高峰晚于雉鹑, 晚活动高峰早于雉鹑, 表现出显著的种间日活动节律差异; 然而, 整合两个维度后, 绿尾虹雉和雉鹑的整体生态位仍然高度重叠, 没有显著分化。本研究表明高山鸡形目物种间的生态位分化体现于多个不同的生态维度, 并且不同物种之间的分化方式有所差异。在空间和时间生态位上的显著分化使雪鹑与同域物种间的竞争压力相对较小, 有利于其实现稳定共存。而绿尾虹雉与雉鹑的整体生态位高度重叠, 建议进一步对其食性开展研究, 探讨营养生态位上的潜在种间分化。     

不同迁移能力和竞争能力的集合种群竞争模式及其模拟

种群竞争是影响生态系统演化的莺要生态过程之一。本文是在徐彩琳和Tilman研究工作的基础上,将竞争系数引入集合种群动力模式,建立了集合种群之间竞争的数学模型,并对5-集合种群的竞争动态进行了计算机模拟研究。结果表明：种群竞争排除与共存受迁移扩散能力和竞争能力影响很大,排除原理在理论上是存在的,在广域集合种群和实际中物种是竞争共存的．共存的条件是其竞争能力与扩散能力呈非线性负相关关系,竞争的结果使物种的强弱序列发生变化。     

科尔沁沙地丘间低地不同生活型物种格局与共存机制

植物种群格局与不同生活型之间物种共存的研究有助于理解物种的生态适应性。丘间低地是科尔沁沙地生态系统发育的一个关键阶段,在此生境中存在多种生活型的植物。采用样线法调查了科尔沁沙地丘间低地100个1 m×1 m样方,根据调查结果选取了分属于4个科3个生活型的15种植物为对象进行格局和共存机制的研究。经格局指数以及Poisson分布的t检验和χ2检验表明:15个物种的种群分布均为聚集分布,但聚集强度差别很大,这可能由种子扩散、物种形态、植物发育时期以及土壤养分条件差异导致;聚集指数DI、CI、m*、GI之间呈极显著正相关关系,表明它们表征共同的生态学意义,相互之间可以替代;多年生草本这一生活型内的物种相互共存较多,多年生草本与其他两个生活型共存较少,这可能与相同生活型物种生态位重叠度大有关。Spearman秩相关和Pearson相关对物种对的检验结果均表明:部分豆科植物与其他科的多年生草本共存机会较大,这可能与豆科植物对改良土壤和减少病虫害有关。物种聚集分布以及不同生活型内几个物种对正相关显著是物种适应环境的结果,对结果显著的部分植物进一步研究可以为沙地植被恢复提供理论指导。     

东太平洋中部中上层鲨鱼群落营养生态位分化

鲨鱼在大洋生态系统中占据着重要的生态地位,其作为顶级捕食者,通过下行效应直接影响生态系统的稳定.稳定同位素技术是目前研究摄食生态学强有力的手段之一,可利用碳氮稳定同位素在食物网中的特性分别指示鲨鱼的食物来源和营养级.本研究选取8种130尾采集自东太平洋中部的中上层鲨鱼,应用稳定同位素绘制其种群生态位图谱,比较不同种群间的生态地位及资源分配方式上的差异.结果表明:不同鲨鱼种群碳、氮稳定同位素比值存在显著差异;8种鲨鱼在东太平洋生态系统中的营养级为4.3~5.4,大青鲨、尖吻鲭鲨与其他6种鲨鱼存在摄食隔离,表现出独特的营养生态地位.这些结果充分证明大洋性中上层鲨鱼并非生态系统的冗余种,其营养生态位的独特性不会被其他捕食者简单地替代和弥补.     

扩散模式对高阶相互作用物种共存机制的影响

物种共存机制是群落生态学研究的核心问题之一,但以成对物种间直接相互作用为主的传统共存理论,并未在实际群落中得到普遍证实。近年来,有研究表明,高阶相互作用,即一个物种对另一个物种的直接作用强度受到其他物种的间接影响,在群落竞争过程中的重要性不断得到发展。目前,对高阶相互作用的理论研究还主要集中在非空间理论模型。事实上,群落中个体的空间分布和扩散模式等对种群动态的影响均至关重要。故考虑空间因素,以三物种为例构建空间显式的群落动态模拟,通过引入不同的物种扩散模式,研究高阶相互作用对群落物种共存结果的影响。研究表明:(1)高阶相互作用可以促进也可能抑制物种共存,具体共存结果取决于高阶相互作用的方向、强度和分类;(2)当全部高阶相互作用都存在,且取值为正时,物种共存位置会发生偏移,原本生态位分化下共存的区域不再共存,而在生态位重叠度较高的区域,物种可以在更大范围的适合度差异下共存;(3)扩散模式对高阶相互作用的上述调节机制有一定的影响,且无论正高阶还是负高阶,当种群趋于局部扩散时,高阶相互作用的正向及负向调节效果均有所减弱。以上结论强调了在理论模型和实际保护工作中考虑相互作用网络的重要性,有助于进一步理解物种共存机制,能够为保护生物多样性提供理论依据。     

基于生态位构建的2-物种集合种群关系动力学分析

结合生态位构建对两局域种群的生存竞争关系的影响,建立了基于生态位构建的2-物种集合种群关系模型,得到了共存平衡点的唯一性和全局稳定性条件,分析了两种群共存过程中空间混沌现象的存在性,并作了实例仿真加以验证．     

基于改进的SURF指数甄选海州湾鱼类群落的关键捕食者

关键捕食者在生态系统中对饵料生物的数量波动、丰富度和空间分布等都起到重要的调控作用。本研究基于在海州湾进行的5个航次的渔业资源底拖网调查资料及胃含物分析数据,通过对传统SURF指数进行改进,甄选海州湾鱼类群落中的关键捕食者。结果表明: 星康吉鳗、长蛇鲻、大泷六线鱼、小眼绿鳍鱼和小黄鱼为海州湾鱼类群落中的关键捕食者,这5种鱼类不但具有较高的连接数,还是多种生物的主要捕食者,在物种连接中具有较强的聚集效应,它们的数量波动会对生态系统能量流动和食物网结构产生较大的影响。通过本方法对关键捕食者进行甄选,不仅考虑了物种间的摄食比例,还将捕捞量及物种的资源量作为重要影响因素,与传统方法相比,具有较大的改进,为关键捕食者的甄选提供了一种新的方法。本研究还发现,物种之间的强相互作用在维护食物网的结构与功能中起着重要作用,加强对关键捕食者的保护,有利于维持生物群落的稳定性和物种多样性,在实施基于生态系统的管理时,要优先保护这些关键物种。星康吉鳗和小黄鱼作为重要的经济鱼种,承受的捕捞压力较高,尤其需要加强保护和管理。     

血雉与其捕食者黄喉貂的时空关系初探

猎物与其捕食者间的种间关系对于维持动物群落的结构与功能至关重要。为了解西南地区中高海拔森林生态系统中较常见的血雉（Ithaginis cruentus）及其捕食者黄喉貂（Martes flavigula）之间的相互作用关系,本研究利用2018至2020年在四川栗子坪国家级自然保护区61个红外相机位点的调查数据（累计相机工作日为13 790 d）,量化分析了两者之间的时空关系。在时间维度上,采用核密度函数绘制了血雉（独立有效照片数n = 156）与黄喉貂（n = 98）的日活动节律曲线,分析结果表明,血雉和黄喉貂在日活动高峰上存在明显的回避现象。在空间维度上,单物种占域模型分析结果显示,影响血雉对位点使用的重要因子是海拔和距水源最近距离,影响黄喉貂占域的生境因子主要是坡度;单季节双物种占域模型的结果显示,（1）在物种作用和环境变量的影响下,黄喉貂存在时血雉对位点的占域率显著低于黄喉貂不存在时;（2）随着海拔的升高,黄喉貂与血雉的空间关系呈现出由分离（物种间的互作因子SIF值小于1）转为重合（物种间的互作因子SIF值大于1）的趋势。本文使用日活动模式和占域模型分析二者在时间生态位和空间生态位上的关系,初步揭示了血雉与其捕食者黄喉貂在时空上呈现不完全分化的特点,为深入理解该地区山地森林生态系统中猎物与其捕食者时空分布关系的研究提供了范例与基础信息。     

植物性别分配与植物群落中物种多样性的维持

植物群落中物种多样性的维持是当代生态学领域中的一个核心问题。根据高斯竞争排除法则,占有相同生态位的物种不能共存。然而植物群落中共存种的数目经常可达数十个以上,难于用以竞争排除法则为基础的竞争共存理论加以解释。本文根据性别分配理论提出了一个新的植物种竞争共存机制;它可以允许占有相同生态位的物种共存。     

长江源区同域分布兔狲、藏狐和赤狐的时空重叠度

兔狲(Otocolobus manul)、藏狐(Vulpes ferrilata)和赤狐(V. vulpes)是青藏高原三江源区域分布的重要小型食肉兽。本研究于2014年6月至2019年9月在青海省长江源区沱沱河和通天河沿岸选取208个位点布设红外相机, 通过所获取的时空分布数据比较了上述3种同域分布小型食肉兽的时空利用情况。通过空间重叠度系数的比较分析, 兔狲和藏狐、兔狲和赤狐以及藏狐和赤狐之间的空间重叠度系数分别为0.25、0.48和0.17, 这表明兔狲、藏狐和赤狐三者在空间利用上存在一定的差异。通过核密度估计方法分析, 兔狲和藏狐属典型的昼行性动物, 而赤狐以夜行性活动为主。兔狲、藏狐和赤狐每个物种在冷暖两季的日活动节律重叠指数分别为0.83、0.78和0.88。两两比较分析表明, 兔狲和藏狐二者的日活动节律重叠指数最高(0.84), 兔狲和赤狐在夜间活动时段存在一定重叠(0.63), 而藏狐和赤狐的时间生态位分化最明显, 重叠指数最低(0.48)。此外, 在暖季, 两两物种之间的日活动节律重叠指数均小于其冷季的重叠指数。综上所述, 长江源区兔狲、藏狐和赤狐3种小型食肉兽可通过空间和时间资源的利用差异来降低物种间的干扰和竞争, 从而达到同域物种共存的目的。     

甘肃祁连山脉雪豹及其同域分布大型食肉动物时间生态位关系

<正>群落内多物种如何共存是群落生态学和生物多样性研究的核心内容之一。经典物种共存理论强调物种之间的生态位分化,侧重于物种对环境的需求,Hutchinson (1957)提出超体积生态位概念,认为物种适合度是由多个因素共同决定,即物种只有在满足其生态位需求的多维空间,     

Predicting the distributions of predator (snow leopard) and prey (blue sheep) under climate change in the Himalaya

Future climate change is likely to affect distributions of species, disrupt biotic interactions, and cause spatial incongruity of predator–prey habitats. Understanding the impacts of future climate change on species distribution will help in the formulation of conservation policies to reduce the risks of future biodiversity losses. Using a species distribution modeling approach by MaxEnt, we modeled current and future distributions of snow leopard (Panthera uncia) and its common prey, blue sheep (Pseudois nayaur), and observed the changes in niche overlap in the Nepal Himalaya. Annual mean temperature is the major climatic factor responsible for the snow leopard and blue sheep distributions in the energy‐deficient environments of high altitudes. Currently, about 15.32% and 15.93% area of the Nepal Himalaya are suitable for snow leopard and blue sheep habitats, respectively. The bioclimatic models show that the current suitable habitats of both snow leopard and blue sheep will be reduced under future climate change. The predicted suitable habitat of the snow leopard is decreased when blue sheep habitats is incorporated in the model. Our climate‐only model shows that only 11.64% (17,190 km²) area of Nepal is suitable for the snow leopard under current climate and the suitable habitat reduces to 5,435 km² (reduced by 24.02%) after incorporating the predicted distribution of blue sheep. The predicted distribution of snow leopard reduces by 14.57% in 2030 and by 21.57% in 2050 when the predicted distribution of blue sheep is included as compared to 1.98% reduction in 2030 and 3.80% reduction in 2050 based on the climate‐only model. It is predicted that future climate may alter the predator–prey spatial interaction inducing a lower degree of overlap and a higher degree of mismatch between snow leopard and blue sheep niches. This suggests increased energetic costs of finding preferred prey for snow leopards – a species already facing energetic constraints due to the limited dietary resources in its alpine habitat. Our findings provide valuable information for extension of protected areas in future.     

Feeding habits and niche partitioning in a predator guild composed of tigers, leopards and dholes in a temperate ecosystem in central Bhutan

We describe the food habits, niche overlap and prey preferences in a predator guild comprised of tigers Panthera tigris , leopards Panthera pardus and dholes Cuon alpinus in a mountainous region of central Bhutan. Scat analyses revealed that these predators consumed 11 different prey species including livestock and rodents, of which leopards consumed 11. The combined relative occurrence of the three species, sambar Cervus unicolor , muntjac Munticus muntjac and wild pig Sus scrofa , constituted 42.7, 33.7 and 71.1% of the tiger, leopard and dhole diets, respectively, while livestock comprised 44.5, 73.4 and 15.9% of the prey consumed, respectively. Regression equations from earlier feeding trials were used to estimate the relative biomass and the numbers of prey consumed. Results showed that sambar featured more frequently than did muntjac and wild pig in the diets of tiger, leopard and dhole and contributed more relative biomass than did muntjac and wild pig. Sambar, muntjac and wild pig together provided 36.9, 28 and 63.1% of the biomass consumed by tigers, leopards and dholes respectively. All else being equal, there was evidence that all three predators ate livestock less than might have been expected on the basis of the abundance and high biomass of this prey category in the area. There was a high dietary niche overlap between the predators (Pianka's overlap index of 0.58–0.92), with a greater overlap between the two felid species than between the felids and the canid. This study provides evidence of a substantial diet overlap among the three sympatric carnivores, and thus highlights the potential for high intra-guild competition among them, especially given the relatively low density of prey.     

祁连山国家公园青海片区雪豹栖息地适宜性评价

雪豹是全球生物多样性保护的旗舰物种,精准评估雪豹栖息地质量可为其种群保护提供科学参考。本文选取地形、气候、植被类型和人为干扰等环境因子,采用景观连接度和MaxEnt模型两种方法,对祁连山国家公园青海片区雪豹栖息地质量进行评价。结果表明：高适宜栖息地主要分布在研究区的西部和中部,适宜面积占青海片区总面积的71.8%～77.5%,两种方法预测的适宜栖息地重叠区为8 979.1 km2,占青海片区总面积的56.7%;39.8%～43.3%的适宜栖息地分布在一般控制区,56.7%～60.2%的适宜栖息地分布在核心保护区;疏勒河源区适宜栖息地面积最多,石羊河源区适宜栖息地面积最少,黑河源区适宜栖息地面积占比最高,约为77.1%～91.8%。适宜栖息地面积整体呈现自东向西递增的趋势。一般控制区分布有较大面积的雪豹栖息地,人类活动将是一般控制区雪豹栖息地的潜在威胁。本研究对优化雪豹就地保护措施提供科学借鉴与参考。     

天山中部雪豹栖息地适宜性研究初报

栖息地丧失和破碎化是许多濒危物种面临的主要威胁,对于分布在高原山地的濒危物种雪豹亦是如此。为了更好地了解天山雪豹的分布情况和栖息地质量,2018年11月至2019年6月,我们在天山中部借助红外相机开展野外调查,调查面积2 425 km²,共获得78个雪豹的出现位点。利用8个物种分布模型对雪豹在此地的生境选择建模,综合结果表明,崎岖度和海拔是影响雪豹分布的主要因素,其中崎岖度大于70和海拔1 700~2 900 m是雪豹出现频率最高的地区。集成模型空间预测显示大部分适宜栖息地集中于乌苏市以东至板房沟分局管辖地区。本研究通过野外调查和模型分析获得了天山中部雪豹的分布区域及环境因子对其的影响,为天山雪豹保护及跨境合作提供科学的数据支撑。     

Snow tracking reveals different foraging patterns of red foxes and pine martens

Red fox (Vulpes vulpes) shares similar prey preferences and co-occurs with several other carnivores, and is together with pine marten (Martes martes), the most common mesocarnivore in the northern boreal forest. Voles are important prey for both species, but it is unclear to what extent they compete for the same food resources in winter. Here, we use 2139 km and 533 km of meticulous snow tracking of red foxes and pine martens to evaluate their food niches. We measured hunting and digging behaviour, whether successful or not, and the effect of snow depth and temperature. Pine martens were restricted to forested habitats, whereas red foxes used a wide range of habitats. Red foxes were found to dig more often than pine martens, 0.67 vs. 0.39 digging events per kilometre. Hunting was less common and similar in both species, about 0.1 hunting event per kilometre. Pine martens were more efficient in hunting and finding food remains compared to red foxes. Increasing snow depth reduced hunting success and also reduced dig success of red foxes. Food niche overlap was small. Red foxes used mostly voles and carrion remains of ungulates, whereas pine martens used cached eggs and small birds. We suggest that caching eggs is an important strategy for pine martens to survive winter in northern latitudes. Snow depth was important for capturing voles, and thick snow cover appeared to mask the effect of vole peaks. Intensified land use, as clear-cutting and leaving slaughter remains from harvest, will benefit red foxes on the expense of pine martens. The ongoing climate change with warmer winters and less snow will likely further benefit the red fox.     

Trophic niche overlap and wild ungulate consumption by red fox and wolf in a mountain area in Italy

Red fox (Vulpes vulpes) and wolf (Canis lupus) are two widespread opportunistic predators living in simpatry in many areas. Nonetheless, scarce information are available on their trophic interactions. We investigated food habits of these two carnivores in a mountain area in Italy and assessed the extent of their trophic niche overlap, focusing on the consumption of wild ungulates. Thereby we analyzed the content of 669 red fox scats and 253 wolf scats collected between May 2008 and April 2009. Red foxes resulted to have a more than three times higher niche breadth than wolves. Vegetables, small mammals, wild ungulates, and invertebrates were major items (altogether 92% of volume) of the red fox annual diet. On the contrary wolf annual diet relied on wild ungulates (94% of volume) with wild boar (Sus scrofa) being the main food item. The degree of trophic niche overlap between the two species was found to be low (Pianka's O = 0.356). Diet variation between the warm and the cold seasons was limited in both species, and higher in red fox than in wolf. The two canids appeared to use wild ungulates unevenly being the former more selective for younger preys, smaller in size (newborn piglets and roe deer Capreolus capreolus fawns), whereas the latter exhibited a preference for medium-sized and large ungulates (10–35 kg wild boar and adult roe deer). Even if wild ungulates represent the main shared food category, the different use of age/weight classes by the two predators, together with their possible consumption as carrions by red fox, suggests a very limited trophic competition between wolf and red fox.This study represents a contribution to the knowledge of trophic interaction in predator–prey systems where sympatric carnivores are present.     

雪豹保护遗传学和基因组学研究进展

雪豹(Panthera uncia)隶属于食肉目猫科豹属,是生活在青藏高原及其周边地区的旗舰物种。随着分子生物学和高通量测序技术的发展,雪豹保护遗传学和保护基因组学研究得到快速的发展,其中非损伤性遗传取样法显著推动了雪豹保护遗传学研究。本文综述了非损伤性遗传取样法在雪豹物种鉴定、个体识别和性别鉴定等研究中的应用,雪豹的系统发生地位、系统地理格局和种群遗传结构及其亚种争议、演化历史、适应性演化和基因组特征等保护遗传学和基因组学方面的研究现状和进展,并对雪豹保护遗传学和基因组学未来发展趋势进行了展望,以期促进雪豹保护生物学研究和保护对策的科学制定。     

Competition versus opportunism: Diet and trophic niche relationship of two sympatric apex carnivores in a tropical forest

There is a little understanding of how apex carnivores partition their diet to coexist. We studied food habit and trophic niche overlap of two apex carnivores, tiger and leopard in the tropical forest of Similipal tiger reserve, eastern India. We used line transect and scat analysis method, to estimate the prey availability and determine the diet and prey selection of two apex carnivores. Tigers consumed mostly large and medium-sized prey, whereas leopards mostly consumed medium and small-sized prey. Both carnivores were not random in their consumption of prey, sambar and wild pigs were selectively consumed by tigers, whereas leopards selectively consumed wild pigs, barking deer and mouse deer. Dietary overlap between two carnivore species was moderate (Pianka's niche overlap index: 0.55), and trophic niche breadth (Levin's standardized niche breadth) of the leopard (0.52) was greater than that of the tiger (0.37). Overall, tigers exhibited specialized feeding habits, whereas leopards showed generalist feeding habits. Our study highlights the opportunistic nature of leopards and probably a reason for the species successfully coexist with tigers.