Effective dispersal rate is a function of habitat size and corridor shape: mechanistic formulation of a two‐patch compartment model for spatially continuous systems

Two‐patch compartment models have been explored to understand the spatial processes that promote species coexistence. However, a phenomenological definition of the inter‐patch ‘dispersal rate’ has limited the quantitative predictability of these models to community dynamics in spatially continuous habitats. Here, we mechanistically rederived a two‐patch Lotka–Volterra competition model for a spatially continuous reaction‐diffusion system where a narrow corridor connects two large habitats. We provide a mathematical formula of the dispersal rate appearing in the two‐patch compartment model as a function of habitat size, corridor shape (ratio of its width to its length), and organism diffusion coefficients. For most reasonable settings, the two‐patch compartment model successfully approximated not only the steady states, but also the transient dynamics of the reaction–diffusion model. Further numerical simulations indicated the general applicability of our formula to other types of community dynamics, e.g. driven by resource‐competition, in spatially homogeneous and heterogeneous environments. Our results suggest that the spatial configuration of habitats plays a central role in community dynamics in space. Furthermore, our new framework will help to improve experimental designs for quantitative test of metacommunity theories and reduce the gaps among modeling, empirical studies, and their application to landscape management.     

Competition between similar invasive species: modeling invasional interference across a landscape

As the number of biological invasions increases, interactions between different invasive species will become increasingly important. Several studies have examined facilitative invader–invader interactions, potentially leading to invasional meltdown. However, if invader interactions are negative, invasional interference may lead to lower invader abundance and spread. To explore this possibility, we develop models of two competing invaders. A landscape simulation model examines the patterns created by two such species invading into the same region. We then apply the model to a case study of Carduus nutans L. and C. acanthoides L., two economically important invasive weeds that exhibit a spatially segregated distribution in central Pennsylvania, USA. The results of these spatially-explicit models are generally consistent with the results of classic Lotka–Volterra competition models, with widespread coexistence predicted if interspecific effects are weaker than intraspecific effects for both species. However, spatial segregation of the two species (with lower net densities and no further spread) may arise, particularly when interspecific competition is stronger than intraspecific competition. A moving area of overlap may result when one species is a superior competitor. In the Carduus system, our model suggests that invasional interference will lead to lower levels of each species when together, but a similar net level of thistle invasion due to the similarity of intra- and interspecific competition. Thus, invasional interference may have important implications for the distribution and management of invasive species.     

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.     

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。     

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.     

Biodiversity and species interactions: extending Lotka–Volterra community theory

A new analysis of the nearly century‐old Lotka–Volterra theory allows us to link species interactions to biodiversity patterns, including: species abundance distributions, estimates of total community size, patterns of community invasibility, and predicted responses to disturbance. Based on a few restrictive assumptions about species interactions, our calculations require only that the community is sufficiently large to allow a mean‐field approximation. We develop this analysis to show how an initial assemblage of species with varying interaction strengths is predicted to sort out into the final community based on the species’ predicted target densities. The sorting process yields predictions of covarying patterns of species abundance, community size, and species interaction strengths. These predictions can be tested using enrichment experiments, examination of latitudinal and productivity gradients, and features of community assembly.     

Coexistence of competing juvenile–adult structured populations

The Leslie-Gower model is a discrete time analog of the competition Lotka–Volterra model and is known to possess the same dynamic scenarios of that famous model. The Leslie–Gower model played a historically significant role in the history of competition theory in its application to classic laboratory experiments of two competing species of flour beetles (carried out by Park in the 1940s–1960s). While these experiments generally supported what became the Competitive Exclusion Principle, Park observed an anomalous coexistence case. Recent literature has discussed Park’s ‘coexistence case’ by means of non-Lotka–Volterra, non-equilibrium dynamics that occur in a high dimensional model with life cycle stages. We study this dynamic possibility in the lowest possible dimension, that is to say, by means of a model involving only two species each with two life cycle stages. We do this by extending the Leslie–Gower model so as to describe the competitive interaction of two species with juvenile and adult classes. We give a complete account of the global dynamics of the resulting model and show that it allows for non-equilibrium competitive coexistence as competition coefficients are increased. We also show that this phenomenon occurs in a general class of models for competing populations structured by juvenile and adult life cycle stages.     

Frequency-dependent community dynamics driven by sexual interactions

Research in community ecology has tended to focus on trophic interactions (e.g., predation, resource competition) as driving forces of community dynamics, and sexual interactions have often been overlooked. Here we discuss how sexual interactions can affect community dynamics, especially focusing on frequency-dependent dynamics of horizontal communities (i.e., communities of competing species in a single ecological guild). By combining mechanistic and phenomenological models of competition, we place sexual reproduction into the framework of modern coexistence theory. First, we review how population dynamics of two species competing for two resources can be represented by the Lotka–Volterra competition model as well as frequency dynamics, and how niche differentiation and overlap produce negative and positive frequency-dependence (i.e., stable coexistence and priority effect), respectively. Then, we explore two situations where sexual interactions change the frequency-dependence in community dynamics: (1) reproductive interference, that is, negative interspecific interactions due to incomplete species recognition in mating trials, can promote positive frequency-dependence and (2) density-dependent intraspecific adaptation load, that is, reduced population growth rates due to adaptation to intraspecific sexual (or social) interactions, produces negative frequency-dependence. We show how reproductive interference and density-dependent intraspecific adaptation load can decrease and increase niche differences in the framework of modern coexistence theory, respectively. Finally, we discuss future empirical and theoretical approaches for studying how sexual interactions and related phenomena (e.g., reproductive interference, intraspecific adaptation load, and sexual dimorphism) driven by sexual selection and conflict can affect community dynamics.     

浙江披山海域主要鱼类时空生态位

依据2015年11月(秋)、2016年2月(冬)、5月(春)、8月(夏)4个航次的渔业资源底拖网调查数据,运用相对重要性指数(IRI)、Shannon指数和Pianka指数对浙江披山海域主要鱼类的时空生态位宽度和重叠进行分析,并结合冗余分析和种间竞争系数研究了主要鱼类间的竞争共存关系及生态位分化。结果表明: 研究期间共采集鉴定鱼类61种,隶属13目29科48属,其中主要鱼类(IRI>100)有19种。绿鳍鱼与丽叶鲹的时间生态位重叠值最高,二者对时间资源利用的同步性最强;孔虾虎鱼与断线舌鳎的空间生态位重叠值和时空生态位重叠值均最高,二者在空间资源序列上的同域性最强,同时,物种对时空二维资源的利用趋于一致。披山海域时空二维生态位显著重叠(Q_ik>0.6)的种对仅占总种对数的5.8%,表明物种时空分布的差异较大,时空生态位重叠受季节变化的影响比较明显,种间竞争系数计测结果与生态位重叠基本一致。冗余分析揭示了主要鱼类与温度、盐度、溶解氧等环境因子之间的关系,进一步解释了物种间的生态位分化。     

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。     

Prey use of wetland benthivorous sunfishes: ontogenetic, interspecific and seasonal variation

The intensity of competitive interactions between fishes is partly determined by prey use and ontogenetic niche shifts. In a wetland where distinct habitat shifts are missing we compared prey use of three generalist benthivorous sunfishes to look for evidence of ontogenetic, interspecific, and “seasonal” variation in prey composition. Diet analysis revealed evidence of diet ontogeny in warmouth (Lepomis gulosus, 30–152 mm standard length, SL), but not in bluespotted sunfish (Enneacanthus gloriosus, 30–47 mm SL) or dollar sunfish (Lepomis marginatus, 30–60 mm SL). Bluespotted and dollar sunfishes consumed small dipteran and amphipod prey and had similar diets in both seasons suggesting a potential for strong interspecific competition. In the dry season, warmouth shifted from using smaller insect prey to larger decapod and fish prey with increasing size. This shift to prey types that were little used by the other species reduced dietary niche overlap with the other sunfishes. After drought and re-flooding (in the wet season), decapods and small fish were less abundant in the wetland and the warmouth ontogenetic shift was less distinct. When matched for gape width, prey composition differed between warmouth and both dollar and bluespotted sunfishes in the wet season, suggesting differences in sunfish foraging modes, but prey use differences were less clear in the dry season when prey were abundant. Both warmouth ontogenetic diet shifts and seasonal variation in prey use (probably mediated by prey abundance) had strong influences on diet overlap and therefore the potential for intra- and interspecific competition between sunfishes in this wetland ecosystem.     

A Lotka–Volterra competition model with seasonal succession

A complete classification for the global dynamics of a Lotka–Volterra two species competition model with seasonal succession is obtained via the stability analysis of equilibria and the theory of monotone dynamical systems. The effects of two death rates in the bad season and the proportion of the good season on the competition outcomes are also discussed.     

阿尔山落叶松主要蛀干害虫的种群空间生态位

对阿尔山落叶松主要蛀干害虫的种群生态位进行了研究,结果表明:在落叶松衰弱立木上,生态位宽度以落叶松八齿小蠹最高,在中龄和老龄树上分别达到0.5782、0.5498,在落叶松中龄和老龄枯死立木上,白带长角天牛的空间生态位宽度分别达到0.5143、0.6294,分布较衰弱木广,而云杉大黑天牛空间生态位宽度在衰弱和枯死立木上都最低。落叶松立木上的空间生态位重叠指数以落叶松八齿小蠹对其他3种优势天牛偏高,天牛之间的空间生态位相似性比例指数均较高且差距较小,说明4种害虫对空间生态位的占据有很大相似程度。4种害虫的空间种间竞争系数均很大,说明对空间和资源的竞争非常激烈,在这种情况下4种害虫能够共存,根本原因是由于取食部位的分化。     

种间竞争的一个新的数学模型—对经典的Lotka—Vol—terra竞争方程的扩充

本文根据营养动力学理论,建立了一类种间竞争的新的数学模型:它是单种群增长的Cui-Lawson模型,在种间竞争上的推广。新的种间竞争模型克服了经典的种间竞争的Lotka-Volteira方程的局限与不足,具有更广泛和复杂的行为,并在特殊条件下以Lotka-Volterra竞争方程为其特例。因此,新的种间竞争的数学模型是更一般的解释性模型,是对经典的Lotka-Voterra竞争方程的扩充。     

北部湾北部生态系统结构与功能——浮游动物空间生态位及其分化

根据2006—2007年北部湾北部海域4个航次调查资料,采用K-优势度曲线法对浮游动物种(类)群进行排序分析,运用Levins公式和Pianka指数分析了主要优势种(类)群的生态位宽度值和生态位重叠程度,并通过除趋势典范对应分析(DCCA)研究生态位分化状况。结果表明,该海区浮游动物优势种(类)群按生态位宽度值可划分为广生态位、中生态位和窄生态位三大类型。其中,广生态位的种(类)群如肥胖软箭虫、亚强次真哲水蚤、长尾类幼体等具有较强的生态适应性,生活范围较广;窄生态位的种(类)群,对环境的适应性较差,对海流和水团等环境因素的变化具有指示意义。生态位宽度能够反映种(类)群丰度季节变化,但不能反映具体丰度大小。北部湾北部全年主要优势种(类)群的生态位宽度值介于0.08—0.77之间,并且各优势种(类)群的生态位宽度差异较大,种对间生态位重叠值介于0.02—0.89之间,平均生态位重叠指数为0.45,种(类)群之间利用资源环境的互补性较强,这与海域内生境多样化并且各优势种对环境适应能力的差异存在一定联系。DCCA分析表明,影响浮游动物分布的主要因子是水深、温度和盐度,其次是叶绿素a和溶解氧。     

Interference competition between an invasive parakeet and native bird species at feeding sites

Interference competition has proved to be a factor of successful establishment of invasive species. This type of competition may have a stronger impact when native species have temporal niche overlap with the invasive species. The ring-necked parakeet Psittacula krameri has been successfully introduced in many countries and its interspecific agonistic behavior has already been reported. The purpose of this study is to analyze the territorial and preemptive interference competition between the ring-necked parakeet and native bird species in a recently colonized area. We used an empirical approach by recording video sequences in gardens equipped with bird feeders in winter. Our results showed that the ring-necked parakeet was the most frequent species at the feeders. Several native species showed temporal niche overlap with the ring-necked parakeet, the highest overlap being with the starling Sturnus vulgaris. The starling was also the species most impacted by interference competition with the parakeet. Our study suggests that, by being most frequently present at the feeders, by demonstrating the most agonistic behavior and by hindering access to food of the other species, the ring-necked parakeet is a superior competitor and may compete with native bird species.     

Ranging Patterns of Two Syntopic Howler Monkey Species (Alouatta guariba and A. caraya) in Northeastern Argentina

To avoid competition, ecologically similar and closely related species tend to differ in their patterns of habitat use when they live in sympatry. We compared ranging patterns of brown howler (Alouatta guariba) and black and gold howler (A. caraya) monkeys living syntopically, i.e., co-occurring and overlapping their ranges in the same habitat within the zone of sympatry, in the Atlantic Forest of northeastern Argentina with the objective of evaluating whether their use of space contributes to the avoidance of interspecific competition for food resources. During 12 mo we collected data on the ranging behavior of 2 groups of each howler species. We analyzed annual and seasonal daily path lengths and movement rates, home range size, use and overlap, habitat and vertical strata use, and intergroup encounters. Black and gold howlers traveled farther and faster during the time of relative food abundance (abundant season) than during the time of relative food shortage (lean season), and their movement rates were affected by group identity and increased with the proportion of fruits in the diet. Brown howlers’ traveling patterns were not affected by any of these factors. Home ranges for both species (95% fixed kernel; brown howlers: 31–70 ha, black-and-gold howlers: 17–112 ha) were among the largest recorded for Alouatta. For both species, core areas (50% fixed kernel) were larger for larger versus smaller groups, and decreased in the lean season compared to the abundant season. Both species showed similar patterns of habitat use, except for a slight vertical stratification. Groups of different species overlapped their ranging areas consistently more and responded to one another less aggressively during encounters than groups of the same species, suggesting that interspecific spatial niche separation for these two syntopic species is not occurring. The vertical stratification, as well as a day-to-day avoidance strategy, may be the only responses of species to one another that could reduce the potentially high levels of competition for food suggested by their elevated trophic niche overlap. A high degree of niche overlap may explain the parapatric distribution of howlers and other closely related and ecologically similar species of primates.     

Fifty years later: trophic ecology and niche overlap of a native and non-indigenous fish species in the western basin of Lake Erie

Since the introduction of white perch (Morone americana) into Lake Erie over 50 years ago, the population size of native yellow perch (Perca flavescens) has decreased up to 79 % and significant changes to the ecosystem have occurred. We examined long-term population estimates and used stable isotopes of carbon (δ ¹³C) and nitrogen (δ ¹⁵N) paired with stomach content analysis to quantify the trophic ecology and niche overlap of adult yellow perch and white perch in the western basin of Lake Erie. We found that changes in yellow perch abundance since 1979 appeared to be better correlated with changes in fishery exploitation rates than with food competition effects from white perch. At the time of this study, yellow perch were found to have higher δ ¹³C values, indicating greater utilization of benthic food resources than white perch, and white perch occupied higher trophic positions based on δ ¹⁵N. The diets of both species varied spatially and seasonally based on stable isotopes and stomach contents, likely driven by changes in prey abundance. Comparison of niche widths using stable isotope population metrics and Schoener diet similarity index suggested a low to moderate degree of niche overlap between species. Isotopic niches of white perch were generally larger than those of yellow perch demonstrating broader resource utilization by this non-indigenous species. We submit that isotopic niche overlap comparisons are more appropriate for studies seeking to understand interactions among populations over course temporal scales, while diet overlap indices, such as the Schoener index provide a means to study fine-scale interactions such as ontogenetic and seasonal diet shifts.     

Increase in ant density promotes dual effects on bee behaviour and plant reproductive performance

Floral rewards do not only attract pollinators, but also herbivores and their predators. Ants are attracted by extrafloral nectaries (EFNs), situated near flowers, and may interfere with the efficiency and behaviour of pollinators. We tested the hypothesis that the impacts of ant–pollinator interactions in plant–pollinator systems are dependent on (1) the seasonal activity of EFNs, which increase ant abundance closer to flowers; (2) consequently, an ant effect, where ants decrease the temporal niche overlap of bees due to predator avoidance; and (3) ant density, where higher densities may negatively affect plant–pollinator interactions and plant performance. We studied two ant–plant–pollinator systems based on Banisteriopsis campestris and Banisteriopsis malifolia plant species. The periods of high ant abundance coincided with plant species blooming. The presence of ants around flowers reduced the visitation rates of the smaller bees and the temporal niche overlap between bee species was not higher than randomly expected when ants had free access. Additionally, we observed variable ant effects on fruit set and duration of bee visits to both Malpighiaceae species when ant density was experimentally kept constant on branches, especially on B. campestris. Our goal was to show the dual role of ant density effects, especially because the different outcomes are not commonly observed in the same plant species. We believe that reduced temporal niche overlap between floral visitors due to ant presence provides an opportunity for smaller bees to improve compatible pollination behaviour. Additionally, we concluded that ant density had variable effects on floral visitor behaviours and plant reproductive performance.