栖息地选择的理论与模型

栖息地选择理论和模型的发展经历了两个主要阶段：理想自由分布模型和空间直观的栖息地选择模型。随着对理论模型假设的放宽，近年来产生了越来越多的新模型。通过对栖息地选择过程的分析，提出了栖息地选择的几个关键环节：栖息地偏好、信息获取、行为决策及选择行为。在建立栖息地选择模型的各个关键环节上均存在大量有待解决的问题。目前对栖息地偏好的研究主要为相关分析，栖息地信息获取的过程仍然是一黑箱；对动物在栖息地选择过程中的行为决策以及对其生理状态的影响尚不了解，而解决这些问题需要生态学、生态学及认知学等多个领域的研究结果支持，也有待新的理论及方法加以充实，甚至还需要其他学科的介入。     

中国大陆鸟类栖息地选择研究十年

栖息地选择研究一直是鸟类生态学研究的重要内容之一。通过对2001年1月至2010年12月10年期间中国大陆鸟类学家在国内外期刊上发表的鸟类栖息地选择研究的论文进行分析,对我国鸟类栖息地选择研究提出展望。10年间,我国鸟类学家共发表有关鸟类栖息地选择或利用的论文170篇,共涉及到鸟类10目31科73种。在这10年里,中国大陆有关鸟类栖息地选择或利用的文章持续增长。然而,我国鸟类栖息地选择的研究也存在着如下问题:(1)存在栖息地选择和栖息地利用误用的现象,这种现象在10年内并无明显改善;(2)在选择研究方法时,很少考虑个体的可获得性、种群密度及抽样尺度对栖息地选择的影响;(3)大多数的栖息地选择的论文没有对鸟类的栖息地选择行为进行研究,也缺乏对其选择的适合度背景去进行研究。根据这些问题,对我国未来的鸟类栖息地选择研究提出了建议。     

动物栖息地选择评估的常用统计方法

利用合适的统计学方法能够更准确地理解动物的栖息地选择。本文通过对2003~2012年期间,10个国际期刊所发表的177篇关于鸟类和兽类栖息地选择论文的30种统计学方法进行分析,简要概述了目前流行的栖息地选择统计学分析方法及特点,同时对同时期的中文文献也进行了简要分析。目前关于动物栖息地选择较为流行的分析方法主要有逻辑斯蒂回归、资源选择函数、成分分析、广义线性模型、多元方差分析、基于欧几里德距离的方法、广义线性混合模型、生态位因子分析、基于个体模型、典型相关分析、物种分布模型等。广义线性模型、逻辑斯蒂回归、多元方差分析和基于欧几里德距离这些方法可以很灵活地用来分析数据,但是缺乏一个有生态学意义的理论框架。资源选择函数和生态位因子分析各自为栖息地选择研究提供了一个统一的理论框架。基于个体的模型是一个自下而上的过程,很难在系统水平形成理论。232篇国内文章中使用较多的方法是主成分分析、Mann-Whitney U检验、t检验、卡方检验、判别分析、方差分析、Vanderloeg选择系数和Scavia选择指数、逻辑斯蒂回归、Kruskal-Wallis H检验和多元回归分析等。在实际研究中,应根据所要解决的研究问题,选择切实可行的分析方法。     

江苏盐城保护区獐的栖息地选择

1999年3月至2000年9月期间在江苏盐城保护区对獐的栖息地选择进行了研究。共记录了123个栖息地利用样方和300个任意样方,在样方中测定了5种生态因子(植被类型、植被高度、植被覆盖度、距水源距离和人为干扰距离)。不同季节獐对植被类型、植被高度和距水源距离的利用有显著差异,对植被覆盖度和人为干扰距离的利用有极显著差异。春季獐偏好选择白茅草滩、大穗结缕草滩、芦苇地和獐毛草滩;夏季主要选择碱蓬地、芦苇地和盐蒿草甸;秋季倾向于选择獐毛草滩、盐蒿草甸、藨草滩和大米草滩。在春、夏、秋季,獐都倾向于选择高度在90-110cm的植被。在不同季节獐都偏好选择植被覆盖度>90%的栖息地,主要选择距离水源200-599m之间的栖息地。     

海南坡鹿恢复生态学研究进展

海南坡鹿的个体总数已经超过1600头,分布地点已经扩展到10多个。海南坡鹿的恢复生态学研究主要包括种群恢复、栖息地管理和行为恢复等3个方面。就种群恢复而言,半野放是一种较稳妥的种群恢复管理方式,应加大投入;野生种群和野放种群的恢复现状缺乏了解,应加强监测与研究。栖息地管理是半野放种群恢复的基础和保障：而圈养、野放和野生种群的栖息地管理工作有待深入。海南坡鹿的恢复也面临行为恢复问题。这方面值得受到更多关注。     

栖息地破碎化与鸟类生存

栖息地破碎化给野生动物带来的不良后果是全球生态学家和保护生物学家共同关心的问题.自从提出栖息地破碎化是导致生物多样性丧失的关键因素之一的论点后,近20年来,栖息地破碎化研究一直是生态学和保护生物学最活跃的前沿研究领域之一.栖息地破碎化是一动态过程,可在多尺度上发生并蕴涵着复杂的空间模式变化.栖息地破碎化对鸟类的生态学效应主要体现在面积效应、隔离效应和边缘效应等.这些效应影响着鸟类的分布、基因交流、种群动态、扩散行为、种间关系和生活史特征等,最终影响着鸟类的生存.介绍和总结了栖息地破碎化过程、研究的理论依据及栖息地破碎化对鸟类生存产生的诸多影响.     

贺兰山蓝马鸡越冬期栖息地的选择

2003年11～12月,在贺兰山采用样带法对蓝马鸡(Crossoptilon auritum)的栖息地选择进行了研究。共测定了25条样带上的62个蓝马鸡栖息地利用样方和50个任意样方的13个生态因子,结果表明,蓝马鸡偏好利用山地针叶林带,避免选择山地草原带、山地疏林草原带、亚高山灌丛和草甸带;偏好利用油松和青海云杉占优势的生境,避免选择山杨、杜松占优势和无树的生境;偏好阳坡和阴坡,避免利用半阳坡和半阴坡。对利用样方和任意样方进行Mann—Whitey U检验,发现利用样方以乔木密度高、灌木密度高、坡度大、隐蔽程度高、雪覆盖浅、高海拔为主要特征。逐步判别分析表明,隐蔽级、灌木密度、乔木密度、距水源距离和坡度具有重要作用,由这5个变量构成的方程对利用样方和任意样方进行正确区分的概率达到91．7％。蓝马鸡的栖息地选择主要与食物条件和隐蔽性有关。     

新疆北塔山雪豹对秋季栖息地的选择

2004年9~10月,对新疆北塔山地区雪豹(Unicaunica)栖息地选择进行了调查。在选定的15条样带上测定了59个利用样方及30个任意样方的6种生境特征(海拔、地形、植被类型、生境平坦度、放牧状况和坡向),其结果如下:Vanderploeg和Scavia选择指数表明:雪豹对海拔、地形、植被类型、生境平坦度和坡向存在选择性:(1)雪豹偏好利用海拔2000~2200m,而避开2600~3000m区域;(2)倾向于悬崖底部和山嵴,避开山坡和山谷;(3)偏好于灌丛,避开森林;(4)避开平坦的开阔地;(5)倾向于选择非放牧区域活动;(6)倾向于选择北坡,避开南坡。对生境特征的主成分分析显示:前3个主成分(海拔、地形和植被类型)的累积贡献率达到75·76%,可以反映雪豹的栖息地特征,同时表明:影响雪豹栖息地选择的主要因素依次为放牧状况、植被类型、地形和生境平坦度。     

甘肃盐池湾黑颈鹤筑巢栖息地偏好及人为干扰的影响

作为高原旗舰物种, 黑颈鹤(Grus nigricolli)是反映高原生态健康状况的重要参考。为了解黑颈鹤如何在多因素作用下适应人类改造过的栖息地环境, 本研究利用遥感解译、最近邻分析与随机森林模型对繁殖于甘肃盐池湾国家级自然保护区党河湿地的黑颈鹤的筑巢栖息地偏好及人为干扰进行研究。2019年和2020年每年的4-9月在党河湿地内对巢位点等数据进行收集。研究结果表明: 党河湿地内巢址与人为干扰的分布位置明显不同, 两者的主要分布区呈现镶嵌状。距深水沼泽距离、距浅水沼泽距离与距湖泊距离是影响黑颈鹤筑巢栖息地选择最关键的3个环境因子。黑颈鹤筑巢时偏好在距离深水沼泽< 125 m、距离浅水沼泽< 130 m、距离湖泊< 270 m的区域内筑巢。黑颈鹤对沼泽、湖泊等资源的偏好是其巢址分布格局的主要驱动力, 而房屋与公路等人为干扰对栖息地选择的影响很小。黑颈鹤筑巢时强烈偏好的栖息地在湿地内占比低、分布聚集且适宜范围有限, 繁殖区域较狭窄。黑颈鹤巢址间距离的上升表明黑颈鹤栖息地质量可能有所下降, 牲畜数量的增长、冬季牧场的多季节利用以及气候变化可能是主要原因。建议在党河湿地内不新增房屋及公路等人为干扰, 同时继续适当限牧, 并给予牧民充足的经济补偿。     

鸟类栖息地片段化研究的理论基础

栖息地片段化是导致许多森林鸟类种群下降的主要原因之一,而对栖息地片段化的形成及其影响的研究已是成为鸟类生态学的研究热点之一。介绍了鸟类栖息地片段化研究的理论基础,即岛屿生物地理学理论、景观生态学理论以及集合种群理论等,并阐述了鸟类栖息地片段化研究范式转变的原因。     

Inferring behavioural mechanisms in habitat selection studies getting the null‐hypothesis right for functional and familiarity responses

Recent studies of animal habitat selection are inferring more detail regarding the behavioural mechanisms involved, like functional responses and familiarity effects. Changes in animals’ use or selection of a habitat type with changing availability are commonly interpreted as a functional response in habitat preference. Studies of familiarity inferred preference for familiar locations from selection for previously visited locations after accounting for habitat‐related preference. We simulated movement paths on discrete landscapes using random walk models with known habitat selection behaviour to assess emergent properties related to habitat selection in the context of functional and familiarity responses. The behavioural interpretation of functional responses relies on the relationship between habitat use and preference. Unfortunately, this relationship is not unique and depends upon the habitat choice mechanism: habitat use can be proportional to preference or proportional to availability (called respectively, hierarchical and simultaneous choice). We found that when the analytical method did not match the choice mechanism, strong functional responses were observed in habitat selection, even though habitat preference was kept constant. Therefore, functional responses need to be discussed in the context of an animal's habitat choice mechanism. In the absence of familiarity‐related preference, we found no familiarity effect while accounting for all habitat variables. However, when habitat models were incomplete (e.g. lack of information about resources and habitats in the landscape) – as in all field studies – spurious preference for familiar locations arose. Our study aids the interpretation of behavioural mechanisms in habitat selection studies, but also calls for a more thorough study of the approaches used to infer behavioural mechanisms in habitat selection studies.     

Experimental evidence for the use of density based interspecific social information in forest birds

Reproductive success and habitat preference are generally assumed to be negatively associated with densities of con- and heterospecific competitors. However, recent theoretical studies have suggested that in some cases habitat preference may have a nonlinear unimodal function in relation to con- or heterospecific competitor densities – intermediate densities being preferred. Such a pattern is expected if con- or heterospecific densities are used as a proximate cue in habitat selection, which may produce benefits by reducing searching costs and providing information about current habitat quality and costs of competition. At low density the use of such cues, and hence habitat selection, are hampered, whereas at high density costs of competition exceed the benefits of using cues, leading to avoidance. Here, we tested this hypothesis by examining whether arboreal migratory birds use the density of resident titmice ( Parus spp.) in habitat selection decisions. Many migrants and titmice species share similar resource needs making titmice density a reliable source of information for migrants. At the scale of habitat patches, we experimentally created a range of titmice densities from low to very high and subsequently measured the density response of migrants. In contrast to the unimodal habitat preference hypothesis, the average species number and total density of migratory birds were positively and linearly correlated with manipulated titmice density. Thus, migrants probably use titmice density as a relative indicator of habitat quality (abundance or quality of food) because foliage gleaners that share similar food resource with titmice, but not ground foragers, showed a positive association with manipulated titmice density. These results emphasize the positive effect of interspecific social information on habitat choice decisions and diversity of migratory bird community.     

动物生境选择研究中的时空尺度

尺度研究已成为生态学上的一个重要概念和研究热点,但是在动物生境选择的研究中尚未引起足够的重视。动物的生境选择包括多层次的判别和一系列等级序位,在各个尺度和水平上具有不同的特征和机制,受到时空尺度的严格限定。繁殖期的时间限制、社群压力、环境变化、动物生理需求的变化决定了动物生境选择的时间制约性;而生境资源的斑块化分布、功能生境之间的相互作用决定了动物生境选择的空间制约性。研究者对时间和空间尺度的选取与应用会直接影响到生境选择研究结果的科学性和实效性。本文从动物生境选择的时空制约性出发,分析了生境选择研究中时间尺度的重要性,叙述了国内外生境选择研究中常见的研究尺度,强调了多尺度研究和长期生态研究的必要性,尺度的选择应该成为生境选择研究的起点和基础。     

Decision-making in a social world: Integrating cognitive ecology and social neuroscience

Understanding animal decision-making involves simultaneously dissecting and reconstructing processes across levels of biological organization, such as behavior, physiology, and brain function, as well as considering the environment in which decisions are made. Over the past few decades, foundational breakthroughs originating from a variety of model systems and disciplines have painted an increasingly comprehensive picture of how individuals sense information, process it, and subsequently modify behavior or states. Still, our understanding of decision-making in social contexts is far from complete and requires integrating novel approaches and perspectives. The fields of social neuroscience and cognitive ecology have approached social decision-making from orthogonal perspectives. The integration of these perspectives (and fields) is critical in developing comprehensive and testable theories of the brain.     

Evolution in fine-grained environments. II. Habitat selection as a homeostatic mechanism

A model of genotype specific habitat selection is developed for an organism subject to within-lifetime environmental fluctuations. Habitat selection is first overlaid upon both hard and soft selection Levene models with either discrete or continuous habitats. It is shown that even if all genotypes have identical physiological and fitness responses within a habitat, habitat selection can still maintain a polymorphism. In other words, physiological divergence is not a necessary prerequisite for divergence in habitat preferences. Within-lifetime environmental variability is then assumed to occur within each chosen habitat. It is shown that habitat selection acts as an evolutionary filter that can enhance the fitness impact of some niches and effectively eliminate the impact of others such that it generally increases the chances for a polymorphism under soft selection. However, density-dependent effects obscure the relationship between physiological fitness and evolutionary outcome. Indeed, it is possible for selection to favor an allele causing its bearers to preferentially go to the niche to which they are least physiologically adapted. Hence, changes in habitat preference can evolve before an organism has completely adapted physiologically to a new habitat. The fitness impact of habitat selection interacts with both homeostatic avoidance mechanisms (i.e., short-term buffering) and with tolerance (long-term) mechanisms. In general, habitat selection will be most favored in those organisms deficient in long-term tolerance. Moreover, habitat selection tends to accentuate selection favoring short-term avoidance mechanisms. Thus, organisms displaying much habitat selection should have poor physiological long-term tolerances but effective physiological short-term avoidance mechanisms. Finally, if the fitness costs associated with habitat selection are too large to be ignored and are comparable for all genotypes, habitat selection directs the selective pressures back onto the physiological homeostatic capabilities. Hence, the very existence and extent of habitat selection depends critically upon the physiological capabilities of the organism.     

Winter microhabitat selection of a threatened pond amphibian in constructed urban wetlands

Mitigating the threat of habitat loss requires actions such as restoring and creating new habitat. In order to effectively achieve this, species habitat requirements and use patterns need to be understood. While many studies have been conducted on the habitat choice of species, these generally focused on habitat use during periods of high activity and detection probability without considering seasonal shifts in habitat use. Understanding habitat selection by frogs during the winter season of low activity may be crucial since it may differ from that used during the summer and may be overlooked as important for population success. We describe the microhabitat use of the threatened green and golden bell frog (Litoria aurea) using radio tracking methods during winter when detection is low and knowledge is limited. We followed 26 individuals between May and July, 2011 to determine whether they selected specific overwintering microhabitats and related this to levels of individual exposure to predators, distance from the edge of the water and temperature of microhabitats. We found that overwintering bell frogs inhabited reeds and rock gabions more frequently than expected and that females used a reduced subset of microhabitats compared to males. Additionally, microhabitats used were more likely to conceal an individual from view, and the majority of overwintering sites were located within 5 m of the edge of the water which may be important for reducing the risk of predation and desiccation. Rock gabions had significantly warmer (1.2°C–1.8°C) mean temperatures than the other microhabitats used. The information presented here can be used in habitat creation and reintroduction programmes to provide habitat which is suitable during both the breeding and non‐breeding season for the conservation of other populations.     

Habitat selection of riparian birds at restoration sites along the Trinity River,California

Riparian habitats in the western United States are imperiled, yet they support the highest bird diversity in arid regions, making them a conservation priority. Riparian restoration efforts can be enhanced by information on species response to variation in habitat features. We examined the habitat selection of four riparian birds known as management indicators at restoration and reference sites along the Trinity River, California. We compared vegetation structure and composition at nest sites, territories, and random points to quantify used versus available habitat from 2012 to 2015. Vegetation in focal species' territories differed between site types, and from available habitat, indicating nonrandom site choice. Birds selected aspects of more structurally complex habitats, such as greater canopy cover, canopy height, and tree species richness. Yellow‐breasted Chats preferred greater shrub cover, and Yellow Warblers preferred greater cover by non‐native Himalayan blackberry. Territory preferences on restoration sites were often a subset of those on reference sites. One exception was canopy height, which was taller on restoration site territories than random points for all species, suggesting that birds preferentially used patches of remnant habitat. Few variables were significant in nest site selection. Restoration plantings along the Trinity River were only 3–10 years old during this study, and have not developed many of the characteristics of mature riparian habitat preferred by birds, but may improve in habitat value over time. Understanding habitat selection is especially important in recently human‐modified environments, where indirect cues used to assess habitat quality may become disassociated from actual habitat quality, potentially creating ecological traps.     

Female choice for genetic complementarity in birds: a review

Data from avian species have played a prominent role in developing and testing theories of female mate choice. One of the most prominent models of sexual selection, the "good genes" model, emphasizes the indirect benefits of female preferences for male ornaments as indicators of a potential sire's additive genetic quality. However, there is growing interest in non-additive sources of genetic quality and mate choice models for self-referential disassortative mating based on optimal levels of genetic dissimilarity. We reviewed the empirical evidence for genetic-complementarity-based female mate choice among birds. We found the evidence for such choice is mixed but in general against the genetic complementarity hypothesis. The lack of evidence for genetic complementarity in many birds may be due to an inability to make the fine distinctions among potential mates based on genes, possibly due to the comparative anosmatic nature of avian sensory system. For some species however there is compelling evidence for genetic complementarity as a criterion used in female mate choice. Understanding the ubiquity of female mate choice based on genetic complementarity and the variation in this source of female preference among and within species remains a challenge.     

Female guppies agree to differ: phenotypic and genetic variation in mate-choice behavior and the consequences for sexual selection

Variation among females in mate choice may influence evolution by sexual selection. The genetic basis of this variation is of interest because the elaboration of mating preferences requires additive genetic variation in these traits. Here we measure the repeatability and heritability of two components of female choosiness (responsiveness and discrimination) and of female preference functions for the multiple ornaments borne by male guppies (Poecilia reticulata). We show that there is significant repeatable variation in both components of choosiness and in some preference functions but not in others. There appear to be several male ornaments that females find uniformly attractive and others for which females differ in preference. One consequence is that there is no universally attractive male phenotype. Only responsiveness shows significant additive genetic variation. Variation in responsiveness appears to mask variation in discrimination and some preference functions and may be the most biologically relevant source of phenotypic and genetic variation in mate-choice behavior. To test the potential evolutionary importance of the phenotypic variation in mate choice that we report, we estimated the opportunity for and the intensity of sexual selection under models of mate choice that excluded and that incorporated individual female variation. We then compared these estimates with estimates based on measured mating success. Incorporating individual variation in mate choice generally did not predict the outcome of sexual selection any better than models that ignored such variation.     

Connectivity, Probabilities and Persistence: Comparing Reserve Selection Strategies

Reserve selection methods are often based on information on species’ occurrence. This can be presence–absence data, or probabilities of occurrence estimated with species distribution models. However, the effect of the choice of distribution model on the outcome of a reserve selection method has been ignored. Here we test a range of species distribution models with three different reserve selection methods. The distribution models had different combinations of variables related to habitat quality and connectivity (which incorporates the effect of spatial habitat configuration on species occurrence). The reserve selection methods included (i) a minimum set approach without spatial considerations; (ii) a clustering reserve selection method; and (iii) a dynamic approach where probabilities of occurrence are re-evaluated according to the spatial pattern of selected sites. The sets of selected reserves were assessed by re-computing species probability of occurrence in reserves using the best probability model and assuming loss of non-selected habitat. The results show that particular choices of distribution model and selection method may lead to reserves that overestimate the achieved target; in other words, species may seem to be represented but the reserve network may actually not be able to support them in the long-term. Instead, the use of models that incorporated connectivity as a variable resulted in the selection of aggregated reserves with higher potential for species long-term persistence. As reserve design aims at the long-term protection of species, it is important to be aware of the uncertainties related to model and method choice and their implications.