Bryophytes and ecological niche theory

Classical niche theory, particularly in terms of competitive exclusion, does not appear to apply as well to bryophytes as to other organisms. Bryophyte communities, as well as those of other plants and of animals, can be thought of in terms of individual species each utilizing particular portions of various resource or habitat continua. Quantitative studies carried out since 1981, particularly those involving niche breadth and overlap, are reviewed. Special attention is given to niche diversification in Sphagnum , Splachnaceae, bryophyte communities in streams, and to ephemeral bryophyte communities. Some bryophyte communities appear to have equilibrium characteristics and to contain species with relatively narrow niche breadths and with no or only partial niche overlap. In many habitats, however, bryophyte communites have non-equilibrium characteristics and diversification of species in microhabitats is opportunistic. Do any bryophyte communities persist long enough for complete saturation by species which have realized niches determined by competitive interactions? Recent studies indicate that this is the case for at least some Sphagnum communities, but that it is the exception not the rule for bryophytes.     

Intraspecific and interspecific competition induces density‐dependent habitat niche shifts in an endangered steppe bird

Interspecific competition is a dominant force in animal communities that induces niche shifts in ecological and evolutionary time. If competition occurs, niche expansion can be expected when the competitor disappears because resources previously inaccessible due to competitive constraints can then be exploited (i.e., ecological release). Here, we aimed to determine the potential effects of interspecific competition between the little bustard (Tetrax tetrax) and the great bustard (Otis tarda) using a multidimensional niche approach with habitat distribution data. We explored whether the degree of niche overlap between the species was a density‐dependent function of interspecific competition. We then looked for evidences of ecological release by comparing measures of niche breadth and position of the little bustard between allopatric and sympatric situations. Furthermore, we evaluated whether niche shifts could depend not only on the presence of great bustard but also on the density of little and great bustards. The habitat niches of these bustard species partially overlapped when co‐occurring, but we found no relationship between degree of overlap and great bustard density. In the presence of the competitor, little bustard's niche was displaced toward increased use of the species' primary habitat. Little bustard's niche breadth decreased proportionally with great bustard density in sympatric sites, in consistence with theory. Overall, our results suggest that density‐dependent variation in little bustard's niche is the outcome of interspecific competition with the great bustard. The use of computational tools like kernel density estimators to obtain multidimensional niches should bring novel insights on how species' ecological niches behave under the effects of interspecific competition in ecological communities.     

Simulated responses to hypothetical fundamental niches

Abstract. Two graphical models of plant competition, the ‘ghost of competition past’ and the ‘hierarchical’ model are compared using a greatly simplified individual-based forest dynamics simulation. Assumed fundamental niche shapes are used to determine the basic growth responses of the species, but competition alters the realized niche. Differences in the two models, amount of niche overlap, disturbance, and removal and invasion of species are examined in simulation experiments. Without disturbance, the realized niche responses reveal abrupt boundaries between species in all cases, and thus the responses are generally platykurtic to rectangular with little overlap. In some cases overlap through the extension of abundance of weaker competitors into the area of greater resources — dominated by better competitors but still within their fundamental niche — creates skewed distributions, as have been observed and simulated in the past. When species are removed or invade, the abrupt boundaries and the lack of difference in final response shape indicate that past removals may be difficult to detect. This difficulty may be important because former species may be influencing the responses that are observed in common non-equilibrium distributions. Even when assumptions favor the illustration of a competitive hierarchy, actual distributions and dynamics do not differentiate between it and a model of the ghost of competition past.     

择伐对思茅松天然林乔木种间与种内关系的影响

通过分析不同择伐强度下思茅松(Pinus kesiya)天然林乔木种群的生态位、种间联结、种内与种间竞争和群落稳定性, 探讨干扰强度对思茅松天然林乔木种群种间与种内关系的影响。结果表明: (1)择伐干扰改变了大多数乔木树种在群落中的优势地位, 思茅松优势地位未受影响, 随着择伐干扰强度的增加, 思茅松在群落中的生态位宽度逐渐变大, 在重度干扰的群落中与红木荷(Schima wallichii)一起成为生态幅度最大的树种。(2)通过种间联结与生态位重叠分析发现, 随着干扰强度的加剧, 乔木种群的种间关系发生变化, 中度干扰群落中物种之间的生态位重叠明显高于未干扰和重度干扰群落, 而在三类群落中, 思茅松与其他种群之间的重叠值较小。随着干扰强度增加, 种群间正联结与负联结的种对所占的比例逐渐增加。(3) Hegyi竞争指数表明, 三类群落中思茅松种群的竞争压力主要来自种间, 而干扰强度进一步增加了思茅松种群的种间竞争强度, 其种内平均竞争指数随着对象木胸径的增加而减少。(4)林窗是思茅松种群种间和种内关系变化的主要原因, 三类群落都处于较不稳定的阶段, 但中度干扰群落的物种丰富度与稳定性都高于未干扰和重度干扰的群落。择伐原则依据森林管理的目的而确定, 如以木材培育为主或森林生态系统服务功能为主。     

How competitive intransitivity and niche overlap affect spatial coexistence

Competitive intransitivity is mostly considered outside the main body of coexistence theories that rely primarily on the role of niche overlap and differentiation. How the interplay of competitive intransitivity and niche overlap jointly affects species coexistence has received little attention. Here, we consider a rock–paper–scissors competition system where interactions between species can represent the full spectra of transitive–intransitive continuum and niche overlap/differentiation under different levels of competition asymmetry. By comparing results from pair approximation that only considers interference competition between neighbouring cells in spatial lattices, with those under the mean-field assumption, we show that 1) species coexistence under transitive competition is only possible at high niche differentiation; 2) in communities with partial or pure intransitive interactions, high levels of niche overlap are not necessary to beget species extinction; and 3) strong spatial clustering can widen the condition for intransitive loops to facilitate species coexistence. The two mechanisms, competitive intransitivity and niche differentiation, can support species persistence and coexistence, either separately or in combination. Finally, the contribution of intransitive loops to species coexistence can be enhanced by strong local spatial correlations, modulated and maximised by moderate competition asymmetry. Our study, therefore, provides a bridge to link intransitive competition to other generic ecological theories of species coexistence.     

基于生态位理论的浙江省旅游城市竞争发展策略

在区域间旅游竞争日趋激烈的背景下,如何明辨竞争对手,采取科学竞争策略,是目前许多旅游城市亟待解决的问题之一。尝试运用生态位重叠理论和生态位态势理论构建了旅游城市竞争关系判断框架,并以浙江省11个旅游城市为研究对象进行了实证分析,发现这11个城市间旅游生态位重叠度都处于较高水平,彼此之间竞争压力比较大。11个旅游城市中,杭州、宁波和温州的旅游综合生态位居前三位,在旅游市场竞争中处于比较有利的地位,而舟山、湖州、衢州分别居倒数前三位,在旅游市场竞争中处于弱势的地位。在此基础上,运用生态位错位、生态位扩充等理论,针对每个城市与其他城市间的生态位重叠度和自身旅游综合生态位的大小,提出了相对应的基本竞争发展策略。     

Pulsed resource availability changes dietary niche breadth and partitioning between generalist rodent consumers

Identifying the mechanisms that structure niche breadth and overlap between species is important for determining how species interact and assessing their functional role in an ecosystem. Without manipulative experiments, assessing the role of foraging ecology and interspecific competition in structuring diet is challenging. Systems with regular pulses of resources act as a natural experiment to investigate the factors that influence the dietary niches of consumers. We used natural pulses of mast‐fruiting of American beech (Fagus grandifolia) to test whether optimal foraging or competition structure the dietary niche breadth and overlap between two congener rodent species (Peromyscus leucopus and P. maniculatus), both of which are generalist consumers. We reconstructed diets seasonally over a 2‐year period using stable isotope analysis (δ¹³C, δ¹⁵N) of hair and of potential dietary items and measured niche dynamics using standard ellipse area calculated within a Bayesian framework. Changes in niche breadth were generally consistent with predictions of optimal foraging theory, with both species consuming more beechnuts (a high‐quality food resource) and having a narrower niche breadth during masting seasons compared to nonmasting seasons when dietary niches expanded and more fungi (a low‐quality food source) were consumed. In contrast, changes in dietary niche overlap were consistent with competition theory, with higher diet overlap during masting seasons than during nonmasting seasons. Overall, dietary niche dynamics were closely tied to beech masting, underscoring that food availability influences competition. Diet plasticity and niche partitioning between the two Peromyscus species may reflect differences in foraging strategies, thereby reducing competition when food availability is low. Such dietary shifts may have important implications for changes in ecosystem function, including the dispersal of fungal spores.     

Community structure, abundance, and morphology

The role of interspecific competition in structuring communities has been a highly debated issue for the last two decades. The nonrandom nature of morphological patterns within communities has been at the center of this controversy. Null models addressing community-wide dispersions in morphology have produced equivocal results and may be based on assumptions that are too restrictive (e.g., competitive exclusion or displacement). If morphological distinctiveness allows species to escape competitive pressures and exhibit higher densities, then a positive relationship should exist between morphological dissimilarity and abundance. We develop a suite of models that evaluates patterns in abundance that are associated with the morphological proximity of a species to other competitors. We evaluated the relationship between morphological distance and abundance from a variety of morphological perspectives, from those representing strictly diffuse interactions to those representing only interactions between a species and its nearest neighbor in morphological space. These models were sufficiently powerful to detect positive associations between abundance and morphological differences in a nocturnal desert rodent guild for which the effects of competition on structure are well established. Models such as these may be more useful than traditional models evaluating morphological dispersions for many reasons. They do not require that communities reach equilibrium before competitive interactions give rise to deterministic structure. They do not suffer from limitations of potentially inaccurate faunal pools or from phylogenetic constraints. Lastly, they may be used as a diagnostic tool in comparative studies to determine the degree to which competitive interactions structure communities.     

Structured environments foster competitor coexistence by manipulating interspecies interfaces

Natural environments, like soils or the mammalian gut, frequently contain microbial consortia competing within a niche, wherein many species contain genetically encoded mechanisms of interspecies competition. Recent computational work suggests that physical structures in the environment can stabilize local competition between species that would otherwise be subject to competitive exclusion under isotropic conditions. Here we employ Lotka-Volterra models to show that interfacial competition localizes to physical structures, stabilizing competitive ecological networks of many species, even with significant differences in the strength of competitive interactions between species. Within a limited range of parameter space, we show that for stable communities the length-scale of physical structure inversely correlates with the width of the distribution of competitive fitness, such that physical environments with finer structure can sustain a broader spectrum of interspecific competition. These results highlight the potentially stabilizing effects of physical structure on microbial communities and lay groundwork for engineering structures that stabilize and/or select for diverse communities of ecological, medical, or industrial utility.     

Competition and niche dynamics from steady-state solutions of dispersal equations

A diffusion model of animal dispersal of the type originated by Skellam (1951, Biometrika 38, 196–218) is used to investigate competitive effects between species. The effects of aggregation, dispersal, and nett growth are related to utility functions specific to each species, and to position on an appropriate axis. Utility, in turn, is defined by whatever underlying competition and resource-use models are being used. The analysis enables niche breadth, overlap, and shift, and different measures of competitive effects, to be explicitly related to underlying competitive models, and the method and meaning of regression-based methods of estimating competitive intensity are clarified. Detailed results are given, for a specific example, concerning the effect of exploratory behaviour, local competitive intensity, and niche separation on equilibrium population numbers, competition coefficients, and on niche overlap and its variation with overall population numbers. Niche overlap is shown to be no indicator of the competition coefficient. The results are shown to accord with observational data.     

Diet breadth,coexistence and rarity in bumblebees

Factors that determine the relative abundance of bumblebee species remain poorly understood, rendering management of rare and declining species difficult. Studies of bumblebee communities in the Americas suggest that there are strong competitive interactions between species with similar length tongues, and that this competition determines the relative abundance of species. In contrast, in Europe it is common to observe several short-tongued species coexisting with little or no evidence for competition shaping community structure. In this study we examine patterns of abundance and distribution in one of the most diverse bumblebee communities in Europe, found in the mountains of southern Poland. We quantify forage use when collecting nectar and pollen for 23 bumblebee species, and examine patterns of co-occurrence and niche overlap to determine whether there is evidence for inter-specific competition. We also test whether rarity can be explained by diet breadth. Up to 16 species were found coexisting within single sites, with species richness peaking in mountain pasture at ~1,000 m altitude. Results concur with previous studies indicating that the majority of pollen collected by bumblebees is from Fabaceae, but that some bee species (e.g. B. ruderatus) are much more heavily dependent on Fabaceae than others (e.g. B. lucorum). Those species that forage primarily on Fabaceae tended to have long tongues. In common with studies in the UK, diet breadth was correlated with abundance: rarer species tended to visit fewer flower species, after correcting for differences in sample size. No evidence was found for similarity in tongue length or dietary overlap influencing the likelihood of co-occurrence of species. However, the most abundant species (which co-occurred at most sites) occupied distinct dietary niche space. While species with tongues of similar length tended, overall, to have higher dietary niche overlap, among the group of abundant short-tongued species that commonly co-occurred there was marked dietary differentiation which may explain their coexistence.     

Niche partitioning in the cestode communities of two elasmobranchs

Several randomization methods have been used to investigate the influence of competitive interactions in shaping parasite community structure. Marine fish parasite communities have often been regarded as unstructured assemblages with little or no resource limitation and, therefore, not prone to competitive influences. In this study, null models were used to assess the niche overlap of cestode communities of two distinct but closely related elasmobranch hosts: the round stingray, Urobatis halleri , Cooper (1863) and the skate, Leucoraja naevus , Muller and Henle (1841). Cestode species distributions were analyzed for randomness with respect to two niche axes: host size class and location within host spiral intestine. Niche overlap of cestode species was calculated for each niche type using MacArthur and Levins' and Pianka's indices, and compared to overlap values obtained from randomly generated communities. Cestodes of both host species were distributed heterogeneously among valves and host size-classes. The majority of parasite species (including all common ones) within U. halleri and all species within L. naevus had significantly nonrandom distributions with respect to at least one niche category. Cestode species pairs tended to overlap significantly less than expected in spiral intestine valves and significantly more than expected among host size classes. We conclude that cestode communities are structured deterministically in a way consistent with expectations based on competition.     

大兴安岭次生林区不同龄级落叶松(Larix gmelinii)生态位特征和竞争关系

生态位能够反映物种在群落中的功能地位,通过生态位可以定量地研究种内、种间、生境三者之间的相互关系。以大兴安岭次生林区优势种群落叶松（Larix gmelinii）为研究对象,运用Levins公式、Pianka指数和Hegyi单木竞争指数模型分析了该优势种的生态位特征、生态位宽度与重叠度以及重叠程度与竞争作用之间的关系。研究结果表明：随落叶松个体的发育其生态位呈先增大后减小的规律性变化;生态位宽度较大的落叶松个体间重叠程度往往也较高,生态位宽度与重叠度呈显著的线性正相关;落叶松个体间的生态位重叠程度与彼此之间的竞争作用无明显相关性、种群内存在互利性生态位重叠现象;生物个体间的竞争作用与研究尺度有关。     

Do plants need niches? Some recent developments in plant community ecology

Species-rich plant communities appear to defy the competitive exclusion principle, showing relatively few obvious niche differences between coexisting species. Here we explore alternatives to the potentially endless search for new niche axes. Spatial aggregation in populations, non-transitive competition, episodes of density-independent mortality and various non-equilibrium theories allow trophically similar species to coexist for extended periods. In perennial plants or annuals with a seed pool, asynchrony between species in recruitment permits coexistence by the 'storage effect'. There is increasing evidence that species-specific herbivores and pathogens regulate populations of tropical trees to low levels at which competitive exclusion does not occur. The wide variety of alternatives to niche differentiation lead us to question whether plants need occupy different niches to coexist.     

乌鲁木齐达坂城山区地衣生态位特征的研究

为揭示达坂城山区地衣对生境资源的利用状况及竞争程度,该研究在野外设置30个样方并调查其物种盖度数据,利用生态位宽度、生态位重叠指数及排序分析,探讨达坂城山区地衣的生态位特征及环境影响因子。结果表明:(1)达坂城山区生态位最宽的为丽黄鳞衣(Rusavskia elegans)和内卷野粮衣(Circinaria contorta),对不同环境具有较强的适应性,其他地衣物种生态位宽度较窄,对环境资源利用程度低。(2)物种间生态位重叠值整体偏低,生态位重叠值较高的物种对极少,地衣物种生态位分化程度高,物种间竞争不激烈。(3)生态位较宽的物种与多数物种之间都存在生态位重叠,但重叠值较低; 一些生态位较窄的物种与其他物种之间的生态位重叠值较高; 生态位重叠和生态位宽度之间并无直接的线性关系。(4)地衣物种沿海拔梯度分布具有差异,海拔、光照强度、湿度、风速和人为干扰是本地区地衣群落物种生态位特征差异的主导环境因子。综上所述,达坂城山区地衣物种由于生境资源的竞争占据各自独特的生态位,形成了生态位的分化,群落较为稳定,沿海拔梯度的生境条件差异是物种差异分布的重要原因。该研究可为地衣群落构建研究提供理论依据,并对该地区地衣物种的多样性和生境保护具有重要意义。     

Evolution in parasite communities

The theoretical aspects of parasite community development are treated under four of the major models on community organization: Nonasymptotic, Asymptotic Equilibrium, Asymptotic Nonequilibrium, and Cospeciation Models. For each model, and for the phases within the Asymptotic Equilibrium Model, the predicted role of evolution in the past and present is described, particularly relating to competition between parasites. The tabulated assumptions of each model enable their identification in real communities, and the answers to five critical questions should provide a means of defining the model appropriate to each system. The study of competition and niche analysis is critical to developments in this field. That very different kinds of parasite communities exist and that different processes operate in each is recognized, but only further study will permit an evaluation of the role of evolution in general.     

Using exclusion rate to unify niche and neutral perspectives on coexistence

The competitive exclusion principle is one of the most influential concepts in ecology. The classical formulation suggests a correlation between competitor species similarity and competition severity, leading to rapid competitive exclusion where species are very similar; yet neutral models show that identical species can persist in competition for long periods. Here, we resolve the conflict by examining two components of similarity – niche overlap and competitive similarity – and modeling the effects of each on exclusion rate (defined as the inverse of time to exclusion). Studying exclusion rate, rather than the traditional focus on binary outcomes (coexistence versus exclusion), allows us to examine classical niche and neutral perspectives using the same currency. High niche overlap speeds exclusion, but high similarity in competitive ability slows it. These predictions are confirmed by a well‐known model of two species competing for two resources. Under ecologically plausible scenarios of correlation between these two factors, the strongest exclusion rates may be among moderately similar species, while very similar and highly dissimilar competitors have very low exclusion rates. Adding even small amounts of demographic stochasticity to the model blurs the line between deterministic and probabilistic coexistence still further. Thus, focusing on exclusion rate, instead of on the binary outcome of coexistence versus exclusion, allows a variety of outcomes to result from competitive interactions. This approach may help explain species coexistence in diverse competitive communities and raises novel issues for future work.     

大型食草动物的聚群规则:印度西部热带干旱森林中有蹄类动物局域和区域尺度的多样性格局

竞争能够塑造自然群落并由其给出可用备择模型检验的一般性聚集规则。关于多种动物集合的竞争结构证据至今还非常少见,我们给出了一个备择模型分析用于检验印度西部热带干旱森林中当地5种大型食草类动物(花鹿Axisaxis,印度大蓝羚Boselaphustragocamelus,水鹿Cervusunicolor,印度瞪羚Gazellabennetti,野猪Susscrofa)是否符合一般性集合规则。使用蒙特卡罗模拟分析了动物的生态位重叠和身体大小比率的类似性,结果表明当地这些种的集合不是竞争结构决定的,其内部阶元结构处于亚最适种丰富度。另外2种有蹄类(印度黑羚Antilopecervicapra,四角羚羊Tetracerusquadricornis)属于地域性种库的组成部分,但缺失于这种集合;身体大小组合在地域性物种集合中不是随机的,然而在当地集合中能够通过随机性机会获得。对于观察到的类型,我们认为由于灭绝留下的统计学空缺似乎最能解释现存集合。由于天敌(大型食肉类)能够减少猎物物种的种间竞争,使得这种无规则生态位配置具有成立的可能性。由于对四角羚羊(T.quadricornis)的生物学和保护现状知之甚少,未将该种包括在该聚集中。我认为对这类较少被人类了解的物种其保护问题被忽视了,今后获得这些稀少的地方特有物种的生态学资料是一项紧急课题.     

An alternative to Lotka-Volterra competition in coarse-grained environments

Competition in a coarse-grained heterogeneous environment is considered. It is assumed that each individual adjust has position within the macrohabitat so as to try to maximize a utility whose value is a function of position within the habitat and of the population density at that position. Robust qualitative conclusions concerning the resulting population behavior relate niche breadth, niche shift, and resource partitioning to changes in population numbers. In particular populations avoid competition by reducing coarse-grained niche overlap so that overlap is a measure not of the degree of competition but of its absence. Specific models of exploitation or interference competition give specific quantitative results which are summarized. Competition equations are derived from underlying models which imply Lotka-Volterra equations in a fine-grained environment (MacArthur, R., and Levins, R. 1967. Amer. Natur.101, 377–385) but which are nonlinear in per capita growth in a coarse-grained one. The findings are relevant to theoretical studies of competition, limiting similarity, species packing, and the evolution of niche position as well as to practical problems of data analysis and resource management. The extension to predator-prey interactions is outlined.     

Niche differentiation in rainforest ant communities across three continents

A central prediction of niche theory is that biotic communities are structured by niche differentiation arising from competition. To date, there have been numerous studies of niche differentiation in local ant communities, but little attention has been given to the macroecology of niche differentiation, including the extent to which particular biomes show distinctive patterns of niche structure across their global ranges. We investigated patterns of niche differentiation and competition in ant communities in tropical rainforests, using different baits reflecting the natural food spectrum. We examined the extent of temporal and dietary niche differentiation and spatial segregation of ant communities at five rainforest sites in the neotropics, paleotropics, and tropical Australia. Despite high niche overlap, we found significant dietary and temporal niche differentiation in every site. However, there was no spatial segregation among foraging ants at the community level, despite strong competition for preferred food resources. Although sucrose, melezitose, and dead insects attracted most ants, some species preferentially foraged on seeds, living insects, or bird feces. Moreover, most sites harbored more diurnal than nocturnal species. Overall niche differentiation was strongest in the least diverse site, possibly due to its lower number of rare species. Both temporal and dietary differentiation thus had strong effects on the ant assemblages, but their relative importance varied markedly among sites. Our analyses show that patterns of niche differentiation in ant communities are highly idiosyncratic even within a biome, such that a mechanistic understanding of the drivers of niche structure in ant communities remains elusive.