Modeling and simulation study of the stoichiometric niche space and niche overlap based on the copula method

Environmental and Ecological Statistics - To qualify niche space and niche overlap, the current methods are based on a strict assumption of the normal distribution and restrict dependence to the...     

人工封育区沙化草地植被生态位研究

采用样线和样方结合法对宁夏人工封育草原进行野外调查,运用重要值、Levins生态宽度指数和Pianka生态位重叠指数对封育外围区、边缘区、核心区3种不同措施下的植物生态位宽度及生态位重叠进行分析,以揭示人工封育区沙化草地植被生态位变化规律。结果表明：在整个人工封育区,重要值之和较大的为黑沙蒿（Artemisia ordosica Krasch.）和刺沙蓬（Salsolaruthenica IIjin.）,分别为689.53、455.83。外围区与边缘区生态位宽度最大的均为黑沙蒿,生态位宽度分别为0.846、0.790,核心区生态位宽度最大的是阿尔泰狗娃花（Heteropappus altaicus（Willd.）Novopokr.）,生态位宽度为0.640,是人工封育区的优势种。生态位重叠计测结果表明,植被的生态位宽度和生态位重叠并不存在直接的线性关系;生态位重叠度平均值和重叠对百分数表现为外围区〉核心区〉边缘区。不同封育措施间生态位宽度和生态位重叠度的差异表明适度封育有利于草地植被恢复,但随着封育时间延长,植物的再生和幼苗的形成会受到一定程度抑制。     

万木林中亚热带常绿阔叶林林隙主要树种的高度生态位

对万木林自然保护区中亚热带常绿阔叶林林隙内外主要树种的生态位宽度和生态位重叠进行了研究。结果表明：桂北木姜子和浙江桂在林隙内外部都占有较高的生态位宽度；生物学特性和生态学特性接近的树种高度生态位重叠较大，反之则较小；在林隙内，出矾和薄叶山矾的高度生态位重叠最大，而桂北木姜子和黄瑞木、尖叶水丝梨，浙江桂和尖叶水丝梨的高度生态位重叠较小；在非林隙内，山矾和黄瑞木的高度生态位重叠最大，桂北木姜子和尖叶水丝梨、黄瑞木的高度生态位重叠较小，表3参13     

On the possible contribution of mixed species flocks to species richness in neotropical avifaunas

Summary Mixed species foraging flocks are a dominant component of the infra-structure of avian communities in neotropical forests. In Amazonia, these flocks consist of pairs of 10–20 species, many of which are permanently associated with mixed flocks. At least half of these flocking species maintain territories that correspond exactly to the flock home range. Small individuals that participate as permanent members of the flocks must adopt the large home range of the larger nucleus species. Therefore, the densities of smaller species are dependent on the availability and density of flocks rather than the availability of food resources. Single pairs of 4 small flocking species with individual body masses of 8 g occupied exclusive territories of 8–12 ha. These were the same exact territories that were defended by at least 6 other flocking species with individual body masses of up to 37 g. Because of their attachment to flocks with large territories, small species are expected to under-utilize available food resources. The under-utilization of food resources is expected to allow smaller species to coexist with greater niche overlap resulting in increased species richness. This hypothesis was tested by quantifying foraging niche in terms of foraging height, foraging maneuver, and prey substrate; and using these values in addition to body mass and bill size (length, depth and width) to determine relative niche overlap between large versus small species pairs.Smaller species had greater foraging overlap than large flocking species and particularly the three smallest species of the genus Myrmotherula; longipennis, axillaris and menetriesii had very high overlap (average foraging niche overlap for the 3 species=0.83±0.12 compared with 0.12±0.19 for all flocking species), similar body sizes (body masses differing by no more then 8%) and similar bill morphologies (maximum ratio in length=1.08, width=1.07, and depth=1.06). These results are consistent with the hypothesis that small species participating in Amazonian mixed flocks can coexist with greater niche overlap because their density is flock dependent rather than resource dependent.     

Individual trophic specialisation and niche segregation explain the contrasting population trends of two sympatric otariids

Individual specialisation is increasingly recognised to be an ecological and evolutionary process having important consequences for population dynamics of vertebrates. The South American fur seal (SAFS) and the South American sea lion (SASL) are two otariid species with similar ecology that coexist in sympatry in the Uruguayan coast. These two species have contrasting trends and widely different population sizes. The underlying reasons for these population trends, unique in their geographical ranges, remain unknown. We studied the foraging ecology of these otariid species over 2 years at the individual- and population levels using the isotopic ratios (δ¹³C–δ¹⁵N) in whiskers of both sexes. We compared the isotope ratios between species and sexes and used several metrics to characterise the degree of overlap and distinctiveness in the use of isotopic niche space at the individual- and population levels. Interspecific trophic niche overlap was minimal, thus ruling out interspecific competition as the cause for the contrasting population trends of both species. At the intraspecific level, both species had sexual segregation in their foraging areas, but each species had a large overlap in the isotopic niches between sexes. While SAFS had a wider niche and generalist individuals, SASL had the narrower niche with a higher degree of individual specialisation. Behavioural constraints during the breeding season, intraspecific competition and a major dependence on resources of the Uruguayan coastal shelf may explain why SASL had a higher trophic individual specialisation and a larger vulnerability in a heavily exploited habitat by fisheries and, by consequence, a locally declining population trend.     

毛竹混交林主要种群多维生态位特征

应用生态位空间分割法，分别考虑资源利用率与否测定毛竹混交林多维资源空间中主要种群多维生态位宽度及多维生态位重叠，结果表明：考虑资源利用率与否对多维生态位宽度的测定有较大的影响，毛竹混交林中主要种群的多维生态位宽度与物种的生物学特性及竞争力密切相关，同一生境中物种的多维生态位重叠可通过生态位分化而减少竞争，研究毛竹混交林主要种群多维生态位特征对毛竹混交林营造具有指导作用．表8参24．     

沙质海岸带木麻黄林与天然林主要种群的生态位特征比较

通过样地调查比较分析了沙质海岸带木麻黄(Casuarina equisetifolia)林与天然林主要种群的生态位宽度和生态位重叠这2种重要的生态位特征。结果表明,依照各种群在2种林分中生态位宽度的变化,可将其划分为生态位扩展型、衰退型、敏感型和稳定型4个种群类型;天然林中的演替优势种在木麻黄林的灌木种群中已有较大数量的分布,且具有较大的生态位宽度值,但由于人工林生态位分化不良,种间竞争相对激烈,群落组成不稳定;天然林中大部分乡土树种生态位宽度较之在木麻黄林中大为增加,但除敏感型种群和潺槁木姜子(Litsea glutinosa)、雀梅藤(Sageretia thea)以外,天然林中其他种群的生态位重叠值较之在木麻黄林中有所降低,说明经过长期的协同进化,天然林种间生态位分化程度高,竞争相对缓和。总体来看,以乡土树种为优势种的天然林生态位配置比人工林更优化,对维持群落的稳定更有意义,因此实现乡土树种与木麻黄的合理配置对建设稳定型沿海防护林体系具有重要的参考价值。     

Interspecific competition between native Eurasian red squirrels and alien grey squirrels: does resource partitioning occur?

In heterogeneous environments, differential niche selection by two competing species will result in niche partitioning so that individuals of each species can maximise their fitness under different sets of environmental variables. Thus, niche partitioning is considered essential to allow co-existence of ecologically related species. To assess whether niche partitioning was occurring between native red squirrels and alien grey squirrels living together in a 13-ha high-quality mixed deciduous woodland in north Italy, we investigated temporal and spatial patterns in their activity and foraging behaviour between 1996 and 1998. We used live trapping and radio-tracking to study numbers, distribution and behaviour of squirrels. Daily and seasonal temporal activity patterns, and activity on the ground and in the trees, were similar in the two species. However, grey squirrels were more tree specialists and had a narrower tree-species niche width than red squirrels, in particular making greater use of oak. Other studies of red and grey squirrels in allopatry show that the two species differ in the extent they utilise oak. Overall, tree-species niche overlap was about 70%. Grey squirrels had larger home ranges than red squirrels. Home ranges and core areas of both species were larger in males than females. Also, intraspecific home range and core-area overlap patterns were similar to those found in allopatric populations of these species. Overall, there was no evidence that the use of space of one species was affected by the other. Our results show that there was no niche partitioning of activity or foraging behaviour in time or space during the study. This suggests that, at moderate grey-squirrel densities, red squirrels are unable to avoid competition with grey squirrels, and that competition for food and/or space will occur when these resources become limiting.     

洞庭湖区退田还湖过程中小型兽类的生态位特征

为了解退田还湖工程实施后的小兽群落状况,2003~2010年对洞庭湖区退田还湖后9种不同类型生境中小型兽类的生态位特征进行了调查.共布放64 432夹日,捕获小型兽3 066只,总的捕获率为4.76%,包括啮齿目10种和食虫目2种.小型兽类各种群的生态位宽度以褐家鼠和黑线姬鼠最高,分别为0.819 8和0.765 6,为广生态位的种类;随后依次是臭鼩、东方田鼠、刺猬、巢鼠、黄胸鼠、小家鼠、黄毛鼠、针毛鼠,分别为0.689 0、0.573 7、0.527 3、0.482 8、0.394 8、0.289 7、0.265 1、0.098 5;社鼠和大足鼠最低,仅在1种生境中有捕获.从生态位宽度指数的季节变化看,黑线姬鼠、褐家鼠、东方田鼠和臭鼩在四季的生态位宽度指数均较高.从空间生态位宽度和生态位重叠指数看,除黑线姬鼠和褐家鼠利用生境较广外,洞庭湖小兽群落可分为3个类群:栖息于环湖丘岗林地生境的社鼠、针毛鼠和黄毛鼠类群,以湖滩生境为主的东方田鼠、巢鼠和臭鼩类群,以及以村庄周围的农田生态系统为主的家鼠类群.从生态位指标和捕获数量看,黑线姬鼠、东方田鼠和褐家鼠是当地需要重点关注的种类.东方田鼠种群数量高峰年份与低谷年份比,其生态位宽度以及与其它种类的生态位重叠指数均较高,高峰年份东方田鼠冬季在双退垸出现,说明退田还湖已扩大了东方田鼠的繁殖基地,对东方田鼠种群发展有利,在种群暴发年份,易形成更加严重的危害.     

生态修复后九曲湾水库浮游动物优势种演替及生态位特征分析

探究生态修复后九曲湾水库浮游动物优势种生态位特征及主要影响因素,为九曲湾水库及相关饮用水源地水库生态系统的保护、修复提供理论和技术支撑。于生态修复前2014年4月(平稳期)、2014年7月(爆发期),2015—2016年生态修复后12月(枯水期)、4月(平水期)、7月(丰水期)进行数据采集,采用优势度、Levins生态位宽度、Petrailis生态位重叠指数等方法,对九曲湾水库浮游动物优势种演替、生态位特征及主要影响因素进行分析。结果表明,九曲湾水库生态修复前共鉴定浮游动物30种,其中轮虫类26种,桡足类4种,枝角类未检出;生态修复后共鉴定浮游动物50种,其中轮虫类30种,枝角类13种,桡足类7种。生态修复前,平稳期到爆发期优势种的更替率为75%,爆发期到生态修复后枯水期优势种的更替率为80%。生态修复后,枯水期到平水期优势种的更替率为66.6%,平水期到丰水期季节更替率为57.14%,共同优势种为裂足臂尾轮虫(Brachionus diversicornis)、汤匙华哲水蚤(Sinocalanus dorrii)。九曲湾水库浮游动物优势种生态位宽度在0.250—0.931之间,生态修复前和生态修复后广生态位种类占全部优势种的75%、58.33%、70%、33%、0。优势种生态位重叠指数在0—0.998之间,生态位重叠程度差异较大,平稳期、爆发期、枯水期、平水期、丰水期生态位重叠指数大于0.6的分别占53.57%、50%、73.33%、66.67%、66.67%。生态修复前影响浮游动物优势种生态位分化的主要因素是水温(WT)、溶解氧(DO),生态修复后影响浮游动物优势种生态位分化的主要因素是总磷(TP)、叶绿素(Chl.a)。研究表明,生态修复前蓝藻水华发生期影响浮游动物生态位分化的环境因子为物理因子,生态修复后影响浮游动物生态位分化的环境因子为营养盐。     

Assessing the likely effectiveness of multispecies management for imperiled desert fishes with niche overlap analysis

A critical decision in species conservation is whether to target individual species or a complex of ecologically similar species. Management of multispecies complexes is likely to be most effective when species share similar distributions, threats, and response to threats. We used niche overlap analysis to assess ecological similarity of 3 sensitive desert fish species currently managed as an ecological complex. We measured the amount of shared distribution of multiple habitat and life history parameters between each pair of species. Habitat use and multiple life history parameters, including maximum body length, spawning temperature, and longevity, differed significantly among the 3 species. The differences in habitat use and life history parameters among the species suggest they are likely to respond differently to similar threats and that most management actions will not benefit all 3 species equally. Habitat restoration, frequency of stream dewatering, non‐native species control, and management efforts in tributaries versus main stem rivers are all likely to impact each of the species differently. Our results demonstrate that niche overlap analysis provides a powerful tool for assessing the likely effectiveness of multispecies versus single‐species conservation plans. Evaluación de la Posible Efectividad del Manejo Multi‐Especie paraPeces de Desierto en Peligro Mediante el Análisis de Traslape de Nichos     

滹沱河流域湿地植被优势种群生态位研究

采用Shannon-Wiener生态位宽度指数和Petraitis生态位重叠指数测定了滹沱河流域湿地植被20个优势种群的生态位宽度和生态位重叠.结果表明,浮叶眼子菜、莎草、藨草等占据较窄的生态位,生态位发生明显特化.苔草、海乳草、黄花蒿、赖草等占据较宽的生态位,生态位泛化比较明显.根据生态位宽度指数将湿地植物分为4个生态类群.在20个优势种群构成的380个种对中,有71个种对间表现出生态位重叠,占整个种对的18.68%.说明优势种群间生态位重叠现象并不普遍,各种群间生态位明显分离,因此不同种群利用资源的方式具有较为显著的差异性.图1表3参15     

Bayesian nonparametric inference for the overlap of daily animal activity patterns

The study of the interaction among species is an active area of research in Ecology. In particular, it is of interest to evaluate the overlap of their ecological niches. Temporal activity is one of the niche’s axes most commonly used to explore ecological segregation among animal species, and many contributions focus on the overlap of this variable. Once the information of the temporal activity is obtained in the wild, the data is treated as a random sample. There exist different methods to estimate the overlap. Specifically, in the case of two species, one possibility is to estimate the density of the temporal activity of each species and then evaluate the overlap between these density functions. This leads naturally to the analysis of circular data. Most of the procedures currently in use impose some rather restrictive assumptions on the probabilistic models used to describe the phenomena, and only provide approximate measures of the uncertainty involved in the process. In this article, we propose a Bayesian nonparametric approach which incorporates a well-defined noninformative prior. We take advantage of the data structure to define such a prior in terms of the predictive distribution. To the best of our knowledge, this is a novel approach. Our procedure is compared with a well-known method using simulated data, and applied to the analysis of real camera-trap data concerning two mammalian species from the El Triunfo biosphere reserve (Chiapas, Mexico).     

Constancy and change in marine predator diets across a shift in oceanographic conditions in the Northern California Current

Variable ocean conditions can greatly impact prey assemblages and predator foraging in marine ecosystems. Our goal was to better understand how a change in ocean conditions influenced dietary niche overlap among a suite of midtrophic-level predators. We examined the diets of three fishes and one seabird off central Oregon during two boreal summer upwelling periods with contrasting El Niño (2010) and La Niña (2011) conditions. We found greater niche specialization during El Niño and increased niche overlap during La Niña in both the nekton and micronekton diet components, especially in the larger, more offshore predators. However, only the two smaller, more nearshore predators exhibited interannual variation in diet composition. Concurrent trawl surveys confirmed that changes in components of predator diets reflected changes in the prey community. Using multiple predators across diverse taxa and life histories provided a comprehensive understanding of food-web dynamics during changing ocean conditions.     

Resource-partitioning and predation impact of a low-latitude myctophid community

This study, based on data collected during summer 1985 in the eastern Gulf of Mexico, examined the degree of overlap in two prime niche parameters, space and food, in 17 of the most abundant myctophid species which inhabit the epipelagic zone at night. Cluster-analyses of vertical distribution information and diet characteristics revealed that while large groups of species overlapped (>60%) in either vertical distributions or diet, when both niche parameters were considered together, little interspecific or intraspecific (size class) overlap occurred. Our data suggest that for myctophids trophic competition is reduced through resource-partitioning, although with considerable overlap at niche boundaries. Niche separation presumably is the result of competition during the evolution of the ecosystem and is maintained presently as diffuse competition: the effect on a species of the combined competition from all other species at that trophic level. We suggest that the large degree of niche overlap enables the packing of over 50 myctophid species in the epipelagic zone at night. Our calculations indicate that myctophid predation nightly removes 2% of the zooplankton biomass. Myctophid predation is selective in that greatest pressure is on certain size classes and types of prey (copepods, ostracods and euphausiids). In the case of copepods, impact is greatest on the larger, more mature stages and hence on the breeding population. It is estimated that myctophids account for at least one-third of the daily production of zooplankton removed from the epipelagic zone by micronekton in the eastern Gulf.     

Can stable isotope ratios provide for community-wide measures of trophic structure?

Stable isotope ratios (typically of carbon and nitrogen) provide one representation of an organism's trophic niche and are widely used to examine aspects of food web structure. Yet stable isotopes have not been applied to quantitatively characterize community-wide aspects of trophic structure (i.e., at the level of an entire food web). We propose quantitative metrics that can be used to this end, drawing on similar approaches from ecomorphology research. For example, the convex hull area occupied by species in delta13C-delta15N niche space is a representation of the total extent of trophic diversity within a food web, whereas mean nearest neighbor distance among all species pairs is a measure of species packing within trophic niche space. To facilitate discussion of opportunities and limitations of the metrics, we provide empirical and conceptual examples drawn from Bahamian tidal creek food webs. These examples illustrate how this methodology can be used to quantify trophic diversity and trophic redundancy in food webs, as well as to link individual species to characteristics of the food web in which they are embedded. Building from extensive applications of stable isotope ratios by ecologists, the community-wide metrics may provide a new perspective on food web structure, function, and dynamics.     

Species-specific tuning increases robustness to sampling bias in models of species distributions: An implementation with Maxent

Various methods exist to model a species’ niche and geographic distribution using environmental data for the study region and occurrence localities documenting the species’ presence (typically from museums and herbaria). In presence-only modelling, geographic sampling bias and small sample sizes represent challenges for many species. Overfitting to the bias and/or noise characteristic of such datasets can seriously compromise model generality and transferability, which are critical to many current applications - including studies of invasive species, the effects of climatic change, and niche evolution. Even when transferability is not necessary, applications to many areas, including conservation biology, macroecology, and zoonotic diseases, require models that are not overfit. We evaluated these issues using a maximum entropy approach (Maxent) for the shrew Cryptotis meridensis, which is endemic to the Cordillera de Mérida in Venezuela. To simulate strong sampling bias, we divided localities into two datasets: those from a portion of the species’ range that has seen high sampling effort (for model calibration) and those from other areas of the species’ range, where less sampling has occurred (for model evaluation). Before modelling, we assessed the climatic values of localities in the two datasets to determine whether any environmental bias accompanies the geographic bias. Then, to identify optimal levels of model complexity (and minimize overfitting), we made models and tuned model settings, comparing performance with that achieved using default settings. We randomly selected localities for model calibration (sets of 5, 10, 15, and 20 localities) and varied the level of model complexity considered (linear versus both linear and quadratic features) and two aspects of the strength of protection against overfitting (regularization). Environmental bias indeed corresponded to the geographic bias between datasets, with differences in median and observed range (minima and/or maxima) for some variables. Model performance varied greatly according to the level of regularization. Intermediate regularization consistently led to the best models, with decreased performance at low and generally at high regularization. Optimal levels of regularization differed between sample-size-dependent and sample-size-independent approaches, but both reached similar levels of maximal performance. In several cases, the optimal regularization value was different from (usually higher than) the default one. Models calibrated with both linear and quadratic features outperformed those made with just linear features. Results were remarkably consistent across the examined sample sizes. Models made with few and biased localities achieved high predictive ability when appropriate regularization was employed and optimal model complexity was identified. Species-specific tuning of model settings can have great benefits over the use of default settings.     

Native ecotypic variation and the role of host identity in the spread of an invasive herbivore, Cactoblastis cactorum

Environmental niche models (ENMs) have gained enormous popularity as tools to investigate potential changes in species distributions resulting from climate change and species introductions. Despite recognition that species interactions can influence the dynamics of invasion spread, most implementations of ENMs focus on abiotic factors as the sole predictors of potential range limits. Implicit in this approach is the assumption that biotic interactions are relatively unimportant, either because of scaling issues, or because fundamental and realized niches are equivalent in a species' native range. When species are introduced into exotic landscapes, changes in biotic interactions relative to the native range can lead to occupation of different regions of niche space and apparent shifts in physiological tolerances. We use an escaped biological control organism, Cactoblastis cactorum (Berg.), to assess the role of the environmental envelope as compared with patterns of host-herbivore associations based on collections made in the native range. Because all nonnative populations are derived from a single C. cactorum ecotype, we hypothesize that biotic interactions associated with this ecotype are driving the species' invasion dynamics. Environmental niche models constructed from known native populations perform poorly in predicting nonnative distributions of this species, except where there is an overlap in niche space. In contrast, genetic isolation in the native range is concordant with the observed pattern of host use, and strong host association has been noted in nonnative landscapes. Our results support the hypothesis that the apparent shift in niche space from the native to the exotic ranges results from a shift in biotic interactions, and demonstrate the importance of considering biotic interactions in assessing the risk of future spread for species whose native range is highly constrained by biotic interactions.     

Sensitivity of species-distribution models to error,bias, and model design: An application to resource selection functions for woodland caribou

Models that predict distribution are now widely used to understand the patterns and processes of plant and animal occurrence as well as to guide conservation and management of rare or threatened species. Application of these methods has led to corresponding studies evaluating the sensitivity of model performance to requisite data and other factors that may lead to imprecise or false inferences. We expand upon these works by providing a relative measure of the sensitivity of model parameters and prediction to common sources of error, bias, and variability. We used a one-at-a-time sample design and GPS location data for woodland caribou (Rangifer tarandus caribou) to assess one common species-distribution model: a resource selection function. Our measures of sensitivity included change in coefficient values, prediction success, and the area of mapped habitats following the systematic introduction of geographic error and bias in occurrence data, thematic misclassification of resource maps, and variation in model design. Results suggested that error, bias and model variation have a large impact on the direct interpretation of coefficients. Prediction success and definition of important habitats were less responsive to the perturbations we introduced to the baseline model. Model coefficients, prediction success, and area of ranked habitats were most sensitive to positional error in species locations followed by sampling bias, misclassification of resources, and variation in model design. We recommend that researchers report, and practitioners consider, levels of error and bias introduced to predictive species-distribution models. Formal sensitivity and uncertainty analyses are the most effective means for evaluating and focusing improvements on input data and considering the range of values possible from imperfect models.