Competing populations on fragmented landscapes with spatially structured heterogeneities: improved landscape generation and mixed dispersal strategies

Interactions between two species competing for space were studied using stochastic spatially explicit lattice-based simulations as well as pair approximations. The two species differed only in their dispersal strategies, which were characterized by the proportion of reproductive effort allocated to long-distance (far) dispersal versus short-distance (near) dispersal to adjacent sites. All population dynamics took place on landscapes with spatially clustered distributions of suitable habitat, described by two parameters specifying the amount and the local spatial autocorrelation of suitable habitat. Whereas previous results indicated that coexistence between pure near and far dispersers was very rare, taking place over only a very small region of the landscape parameter space, when mixed strategies are allowed, multiple strategies can coexist over a much wider variety of landscapes. On such spatially structured landscapes, the populations can partition the habitat according to local conditions, with one species using pure near dispersal to exploit large contiguous patches of suitable habitat, and another species using mixed dispersal to colonize isolated smaller patches (via far dispersal) and then rapidly exploit those patches (via near dispersal). An improved mean-field approximation which incorporates the spatially clustered habitat distribution is developed for modeling a single species on these landscapes, along with an improved Monte Carlo algorithm for generating spatially clustered habitat distributions.      

Coexistence under positive frequency dependence

Negative frequency dependence resulting from interspecific interactions is considered a driving force in allowing the coexistence of competitors. While interactions between species and genotypes can also result in positive frequency dependence, positive frequency dependence has usually been credited with hastening the extinction of rare types and is not thought to contribute to coexistence. In the present paper, we develop a stochastic cellular automata model that allows us to vary the scale of frequency dependence and the scale of dispersal. The results of this model indicate that positive frequency dependence will allow the coexistence of two species at a greater rate than would be expected from chance. This coexistence arises from the generation of banding patterns that will be stable over long time-periods. As a result, we found that positive frequency-dependent interactions over local spatial scales promote coexistence over neutral interactions. This result was robust to variation in boundary conditions within the simulation and to variation in levels of disturbance. Under all conditions, coexistence is enhanced as the strength of positive frequency-dependent interactions is increased.     

Spatial mosaic formation through frequency-dependent selection in Müllerian mimicry complexes

Although contemporary models of Müllerian mimicry have considered the movement of interfacial boundaries between two distinct mimetic forms, and even the possibility of polymorphisms in two patch systems, no model has considered how multiple forms of Müllerian mimics might evolve and be maintained over large geographical areas. A spatially explicit individual-based model for the evolution of Müllerian mimicry is presented, in which two unpalatable species are distributed over discrete cells within a regular lattice. Populations in each cell are capable of genetic drift and experience localized dispersal as well as frequency-dependent selection by predators. When each unpalatable prey species was introduced into a random cell and allowed to spread, then mimicry evolved throughout the system in the form of a spatial mosaic of phenotypes, separated by narrow "hybrid zones". The primary mechanism generating phenotypic diversity was the occasional establishment of new mutant forms in unoccupied cells and their subsequent maintenance (and spread) through frequency-dependent selection. The mean number of discrete clusters of the same morph that formed in the lattice was higher the higher the intensity of predation, and higher the lower the dispersal rate of unpalatable prey. Under certain conditions the hybrid zones moved, in a direction dependent on the curvature of their interfacial boundaries. However, the mimetic mosaics were highly stable when the intensity of predation was high and the rate of prey dispersal was low. Overall, this model highlights how a stable mosaic of different mimetic forms can evolve from a range of starting conditions through a combination of chance effects and localized frequency-dependent selection.     

Movement behaviour determines competitive outcome and spread rates in strongly heterogeneous landscapes

Classical models for biological invasions were single-species models in homogeneous landscapes, but most invasions happen in the presence of interacting species and in heterogeneous environments. The combination of spatial variation and species interaction could alter the spreading process significantly. For example, the ‘environmental heterogeneity hypothesis of invasions’ posits that heterogeneity offers more opportunities for invaders and reduces the negative impact on native species. Environmental heterogeneity offers an obvious coexistence mechanism on the regional scale if two or more competing species have different spatial niches, i.e. if the local competitive advantage changes in space. We consider a more subtle mechanism of space use through individual movement behaviour when the local competitive advantage remains with the same species. Specifically, we model the densities of two species, diffusing and competing in an infinite landscape consisting of two types of patches. We include individual behaviour in terms of movement rate and patch preference. We consider the scenario that one of the species is the stronger local competitor in both patch types. We then uncover a number of mechanisms—based solely on movement behaviour—through which these two species can coexist regionally, how the inferior competitor can replace the superior competitor globally, or how a bistable situation can arise between the two. We calculate mutual invasion conditions as well as mutual spatial spread rates, and we show that spread rates may depend on movement parameters in unexpected ways.     

Competition and coexistence in regional habitats

Habitat heterogeneity plays a key role in the dynamics and structures of communities. In this article, a two-species metapopulation model that includes local competitive dynamics is analyzed to study the population dynamics of two competing species in spatially structured habitats. When local stochastic extinction can be ignored, there are, as in Lotka-Volterra equations, four outcomes of interspecific competition in this model. The outcomes of competition depend on the competitive intensity between the competing pairs. An inferior competitor and a superior competitor, or two strongly competing species, can never stably coexist, whereas two weak competitors (even if they are very similar species) may coexist over the long term in such environments. Local stochastic extinction may greatly affect the outcomes of interspecific competition. Two competing species can or cannot stably coexist depending not only on the competitive intensity between the competing pairs but also on their precompetitive distributions. Two weak competitors that have similar precompetitive distributions can always regionally coexist. Two strongly competing species that competitively exclude each other in more stable habitats may be able to stably coexist in highly heterogenous environments if they have similar precompetitive distributions. There is also a chance for an inferior competitor to coexist regionally or even to exclude a superior competitor when the superior competitor has a narrow precompetitive distribution and the inferior competitor has a wide precompetitive distribution.     

Coexistence of cycling and dispersing consumer species: Armstrong and McGehee in space

Two competing consumer species may coexist using a single homogeneous resource when the more efficient consumer--the one having the lowest equilibrium resource density--has a more nonlinear functional response that generates consumer-resource cycles. We extend this model of nonequilibrium coexistence, as proposed by Armstrong and McGehee, by putting the interaction into a spatial context using two frameworks: a spatially explicit individual-based model and a spatially implicit metapopulation model. We find that Armstrong and McGehee's mechanism of coexistence can operate in a spatial context. However, individual-based simulations suggest that decreased dispersal restricts coexistence in most cases, whereas differential equation models of metapopulations suggest that a low rate of dispersal between subpopulations often increases the coexistence region. This difference arises in part because of two potentially opposing effects on coexistence due to the asynchrony in the temporal dynamics at different locations. Asynchrony implies that the less efficient species is more likely to be favored in some spatial locations at any given time, which broadens the conditions for coexistence. On the other hand, asynchrony and dispersal can also reduce the amplitude of local population cycles, which restricts coexistence. The relative influence of these two effects depends on details of the population dynamics and the representation of space. Our results also demonstrate that coexistence via the Armstrong-McGehee mechanism can occur even when there is little variation in the global densities of either the consumers or the resource, suggesting that empirical studies of the mechanisms should measure densities on several spatial scales.     

Taxonomic relatedness and spatial structure of a shelf benthic gastropod assemblage

Aim Phylogenetically related species share attributes that lead to common responses to environmental conditions, but which could also produce the exclusion of species by its relatives. These processes could generate the patterns of phylogenetic attraction or repulsion in local communities, where related species would tend to coexist more or less than expected by chance. This paper aims to (1) analyse the phylogenetic structure of a benthic gastropod assemblage in the south‐western Atlantic Ocean (SAO); (2) explore the linkages between phylogenetic structure and spatial distribution patterns; (3) compare outcomes driven by the analysis of presence‐only data and predictive species distribution models; and (4) explore which aspects of the gained knowledge can be useful to the design of sound conservation and/or management actions. Location Uruguayan shelf and slope in the SAO. Methods Spatial patterns in taxonomical relatedness were assessed using (1) raw presence/absence data (i.e. realized niche approach) and (2) reconstruction of the potential composition of the assemblage from niche modelling (i.e. fundamental niche approach). Null models were used to test hypotheses on assemblage structure. Results Significant departures from the null hypothesis that all species were drawn from the same assemblage were observed irrespectively of the approach, indicating the existence of non‐random structures. However, a high proportion of local communities can be thought as random subsets of the regional species pool. This lack of a strong signal of a taxonomic effect could be related to the absence of a linkage between taxonomic distances and ecological similarities. Main conclusions Our results suggest a random assembly of local communities from the regional species pool and/or niche filtering independent of phylogeny as main determinants of local community composition. We also suggest that local assemblages displaying significantly higher (or lower) than expected taxonomic relatedness should be taken into consideration for designing spatially explicit conservation measures.     

The coexistence of hybrid and parental Daphnia: the role of parasites

Parasite driven time-lagged negative frequency-dependent selection of hosts has been studied in natural populations by following changes in host genotype frequencies over time. However, such dynamics have not been considered at higher taxonomic levels, for example, between parental species and their hybrids. In a field study on a Daphnia hybrid system, we observed that one Daphnia taxon first was relatively under-infected, but became over-infected after a strong increase in frequency. This finding is consistent with the idea of parasite evolution towards the most frequent host taxon. In two experiments, we investigated whether the assumptions made by a model of negative frequency-dependent selection apply to our host taxa system. First, we showed that the parasite can change the outcome of taxa competition and secondly, we confirmed that the over-infection of one host taxon observed in the field has a genetic basis. Our results indicate that the incorporation of host-parasite interactions at the species level may allow us to gain a more complete picture of forces driving dynamic taxa coexistence in Daphnia hybrid systems. More generally, we suggest that if hybrids coexist in sympatry with parental taxa, the infection patterns as observed under natural conditions may be rather temporal and unstable.     

Coexistence through mutualist‐dependent reversal of competitive hierarchies

Mechanisms that allow for the coexistence of two competing species that share a trophic level can be broadly divided into those that prevent competitive exclusion of one species within a local area, and those that allow for coexistence only at a regional level. While the presence of aphid‐tending ants can change the distribution of aphids among host plants, the role of mutualistic ants has not been fully explored to understand coexistence of multiple aphid species in a community. The tansy plant (Tanacetum vulgare) hosts three common and specialized aphid species, with only one being tended by ants. Often, these aphids species will not coexist on the same plant but will coexist across multiple plant hosts in a field. In this study, we aim to understand how interactions with mutualistic ants and predators affect the coexistence of multiple species of aphid herbivores on tansy. We show that the presence of ants drives community assembly at the level of individual plant, that is, the local community, by favoring one ant‐tended species, Metopeurum fuscoviride, while preying on the untended Macrosiphoniella tanacetaria and, to a lesser extent, Uroleucon tanaceti. Competitive hierarchies without ants were very different from those with ants. At the regional level, multiple tansy plants provide a habitat across which all aphid species can coexist at the larger spatial scale, while being competitively excluded at the local scale. In this case, ant mutualist‐dependent reversal of the competitive hierarchy can drive community dynamics in a plant–aphid system.     

Reproductive interference hampers species coexistence despite conspecific sperm precedence

Negative interspecific mating interactions, known as reproductive interference, can hamper species coexistence in a local patch and promote niche partitioning or geographical segregation of closely related species. Conspecific sperm precedence (CSP), which occurs when females that have mated with both conspecific and heterospecific males preferentially use conspecific sperm for fertilization, might contribute to species coexistence by mitigating the costs of interspecific mating and hybridization. We discussed whether two species exhibiting CSP can coexist in a local environment in the presence of reproductive interference. First, using a behaviorally explicit mathematical model, we demonstrated that two species characterized by negative mating interactions are unlikely to coexist because the costs of reproductive interference, such as loss of mating opportunity with conspecific partners, are inevitably incurred when individuals of both species are present. Second, we experimentally examined differences in mating activity and preference in two Harmonia ladybird species known to exhibit CSP. These behavioral differences may lead to local extinction of H. yedoensis because of reproductive interference by H. axyridis. This prediction is consistent with field observations that H. axyridis uses various food sources and habitats whereas H. yedoensis is confined to a less preferred prey item and a pine tree habitat. Finally, by a comparative approach, we observed that niche partitioning or parapatric distribution, but not sympatric coexistence in the same habitat, is maintained between species with CSP belonging to a wide range of taxa, including vertebrates and invertebrates living in aquatic or terrestrial environments. Taken together, it is possible that reproductive interference may destabilize local coexistence even in closely related species that exhibit CSP.     

花柱卷曲性的维持及功能性别特化

花柱多态型的型比通过选型交配形成的负频率依赖的选择来维持。种群空间格局对于频率依赖选择所维持的花柱多态型是至关重要的。该研究使用摆放实验构建不同型比的马来良姜(Alpinia mutica)人工种群, 以评价花柱卷曲性促进选型交配的功能。使用邻域模型调查了云南草蔻(Alpinia blepharaocalyx)种群中的目标植株的坐果数、结籽数及其邻域中花序的数量和型偏以探讨在局域尺度上目标植株的雌性适合度是否受负频率依赖的选择。通过比较云南草蔻上举型和下垂型各繁殖性状的差异以及两型的功能性别的计算以评估两型功能性别可能存在的特化趋势。摆放实验结果说明花柱卷曲性形成的二型种群可以促进型间花粉传递。在局域尺度上, 云南草蔻两型倾向于同型聚集分布, 这一分布特点反映出其克隆生长特性。统计结果说明: 邻域中异型花粉供体相对丰富度对两型目标植株都无影响; 目标植株雌性适合度在邻域中不受负频率依赖选择的影响。这一结果并不支持负频率依赖的选择在该研究尺度上是影响种群型比结构的主要机制。云南草蔻两型个体在花期和果期的繁殖指标上不存在显著差异; 功能性别计算结果显示选型交配是种群内唯一的交配模式时, 下垂型功能性别偏雌性而上举型功能性别偏雄性。这一功能性别特化的趋势可能是通过对上举型的雄性功能的选择而造成的, 并可能与种群型结构相关。当功能性别计算只考虑自交而忽略近交衰退时, 种群功能性别偏向雌性。     

入侵杂草化感作用的细胞自动机模拟研究

细胞自动机模型(Cellular Automata Model,简称CA模型)是一种能够表现系统复杂行为的模拟方法,适于研究植物群落时空动态过程.本文利用CA模型,模拟具有化感作用的外来种入侵原有物种所构成植被的过程.模型由产生化感物质的外来种和两个对化感物质敏感性不同的本地种组合成不同类型的群落,利用化感物质作用下受体物种生物活性响应模型及种子扩散负指数分布模型,模拟外来杂草和本地种分布格局的时空动态变化.结果表明,外来种可成功地完全入侵由两个对化感物质敏感的本地种构成的群落空间,但对于由对化感物质敏感的一个本地种及对化感物质具有抗性的另一个本地种构成的群落,外来种只能够与本地种共存.     

Biome transitions as centres of diversity: habitat heterogeneity and diversity patterns of West African bat assemblages across spatial scales

It is widely accepted that species diversity is contingent upon the spatial scale used to analyze patterns and processes. Recent studies using coarse sampling grains over large extents have contributed much to our understanding of factors driving global diversity patterns. This advance is largely unmatched on the level of local to landscape scales despite being critical for our understanding of functional relationships across spatial scales. In our study on West African bat assemblages we employed a spatially explicit and nested design covering local to regional scales. Specifically, we analyzed diversity patterns in two contrasting, largely undisturbed landscapes, comprising a rainforest area and a forest‐savanna mosaic in Ivory Coast, West Africa. We employed additive partitioning, rarefaction, and species richness estimation to show that bat diversity increased significantly with habitat heterogeneity on the landscape scale through the effects of beta diversity. Within the extent of our study areas, habitat type rather than geographic distance explained assemblage composition across spatial scales. Null models showed structure of functional groups to be partly filtered on local scales through the effects of vegetation density while on the landscape scale both assemblages represented random draws from regional species pools. We present a mixture model that combines the effects of habitat heterogeneity and complexity on species richness along a biome transect, predicting a unimodal rather than a monotonic relationship with environmental variables related to water. The bat assemblages of our study by far exceed previous figures of species richness in Africa, and refute the notion of low species richness of Afrotropical bat assemblages, which appears to be based largely on sampling biases. Biome transitions should receive increased attention in conservation strategies aiming at the maintenance of ecological and evolutionary processes.     

Dispersal-mediated coexistence of competing predators

Models of metapopulations have often ignored local community dynamics and spatial heterogeneity among patches. However, persistence of a community as a whole depends both on the local interactions and the rates of dispersal between patches. We study a mathematical model of a metacommunity with two consumers exploiting a resource in a habitat of two different patches. They are the exploitative competitors or the competing predators indirectly competing through depletion of the shared resource. We show that they can potentially coexist, even if one species is sufficiently inferior to be driven extinct in both patches in isolation, when these patches are connected through diffusive dispersal. Thus, dispersal can mediate coexistence of competitors, even if both patches are local sinks for one species because of the interactions with the other species. The spatial asynchrony and the competition-colonization trade-off are usual mechanisms to facilitate regional coexistence. However, in our case, two consumers can coexist either in synchronous oscillation between patches or in equilibrium. The higher dispersal rate of the superior prompts rather than suppresses the inferior. Since differences in the carrying capacity between two patches generate flows from the more productive patch to the less productive, loss of the superior by emigration relaxes competition in the former, and depletion of the resource by subsidized consumers decouples the local community in the latter.     

Competition and coexistence in spatially subdivided habitats

While non-spatial models predict that like species cannot stably coexist, empirical studies suggest that similar species have similar distributions due to shared habitat requirements. A model is developed to discuss competition and coexistence in subdivided but locally stable habitats. The model predicts that in some cases it is possible for one species to exclude the other species from a geographic region, while in other cases two competing species can stably coexist. The equilibrium level and the fraction of doubly occupied patches, if there is coexistence, are determined by the strength of competition on colonization and exclusion in such a system. Also, it is possible for two ecologically identical species to stably coexist, and two asymmetrically competing species can coexist when there is a trade-off between local competition ability and invasion ability. When rescue effects are considered, the stable region at internal equilibrium point would be reduced, but the fraction of doubly occupied patches would be enlarged.     

The coherence problem with the Unified Neutral Theory of Biodiversity

The Unified Neutral Theory of Biodiversity (UNTB), proposed as an alternative to niche theory, has been viewed as a theory that species coexist without niche differences, without fitness differences, or with equal probability of success. Support is claimed when models lacking species differences predict highly aggregated metrics, such as species abundance distributions (SADs) or species area distributions (SARs). Here, I summarize why UNTB generates confusion, and is not actually relevant to niche theory (i.e. an explanation for why and how many species coexist). Equal probability is not a theory, but lack of one; it does not include or exclude any process relevant to coexistence of competitors. Models lacking explicit species can make useful predictions, but this does not support neutral theory. I provide s suggestions that could help reduce confusion generated by the debate.     

Grazed Pannonian grassland beta-diversity changes due to C4 yellow bluestem

This study investigates how yellow bluestem affects biodiversity in a typical Pannonian grassland. Beta diversity (i.e. the finescale spatial variability of species compositions), was estimated by the realized number of species combinations sampled at various scales. Sampling was performed by a standard protocol. Presences of plant species were recorded along 52.2 m long belt transect of 1044 units of 0.05x0.05 m contiguous microquadrats. According to the results the massive presence of tested C₄ grass significantly reduced species richness of the grassland. Beta diversity assessment revealed that 90% of species combinations were lost due to yellow bluestem invasion. Fine-scale spatial pattern analyses showed complete local extinctions of other species from microsites dominated by yellow bluestem. This local extinction is enhanced by the specific clonal architecture of this species and by the accumulation of litter. Other dominant grasses had no effect on fine scale diversity, i.e. they could coexist well with other elements of the local flora. This study presents currently developed microhabitat types, forecasts and also draws attention to the danger that climate warming will probably enhance the spread of this detrimental C₄ species.     

Opposite shell-coiling morphs of the tropical land snail Amphidromus martensi show no spatial-scale effects

Much can be learned about evolution from the identification of those factors maintaining polymorphisms in natural populations. One polymorphism that is only partially understood occurs in land snail species where individuals may coil clockwise or anti-clockwise. Theory shows that polymorphism in coiling direction should not persist yet species in several unrelated groups of land snails occur in stably polymorphic populations. A solution to this paradox may advance our understanding of evolution in general. Here, we examine two possible explanations: firstly, negative frequency-dependent selection due to predation; secondly, random fixation of alternative coiling morphs in tree-sized demes, giving the impression of wider polymorphism. We test these hypotheses by investigating morph-clustering of empty shells at two spatial scales in Amphidromus martensi populations in northern Borneo: the spatial structure of snail populations is relatively easy to estimate and this information may support one or other of the hypotheses under test. For the smaller scale we make novel use of a statistic previously used in botanical studies (the K-function statistic), which allows clustering of more than one morph to be simultaneously investigated at a range of scales and which we have corrected for anisotropy. We believe this method could be of more general use to ecologists. The results show that consistent clustering or separation of morphs cannot be clearly detected at any spatial scale and that predation is not frequency-dependent. Alternative explanations that do not require strong spatial structuring of the population may be needed, for instance ones involving a mechanism of selection actively maintaining the polymorphism.     

Spatial variation in the fitness of divergent aposematic phenotypes of the poison frog, Dendrobates tinctorius

Aposematic species use brightly coloured signals to warn potential predators of their unpalatability. The function of these signals is largely believed to be frequency-dependent. All else being equal, stabilizing selection is expected to constrain the evolution of novel signals. However, despite the expected frequency-dependent function of aposematic signals, interpopulation variation in aposematic signals is ubiquitous in nature. Here, we used clay models of the poison frog Dendrobates tinctorius to test the nature of selection in regions containing varying frequencies of frogs possessing the local aposematic signal. Our findings support a role for stabilizing selection in maintaining the local signal type in a region of high signal frequency; however, we observe a lack of stabilizing selection at one site coincident with a decrease in the density of frogs possessing the local signal. Spatial variation in local aposematic signal frequencies may facilitate the evolution of novel signal types by altering the adaptive landscape for divergent aposematic phenotypes. Our results provide evidence for spatial variation in the selective regime acting on aposematic signals within an established aposematic system and highlight the need for further study of the nature of selection acting across different spatial scales in diverse aposematic systems.     

Spatial processes that maintain biodiversity in plant communities

The composition of communities of sessile organisms, and the change in species diversity with time, is a spatially explicit phenomenon. Three spatial factors clearly affect diversity: (1) the structure and heterogeneity of the landscape that limits species immigration and ultimate community size; (2) neighborhood interactions that determine colonization and extinction rates and influence residence times of local populations; and (3) disturbances that open spatially contiguous areas for recolonization by less abundant species. The importance of these three factors was first reviewed and then examined with a spatially explicit, multi-species model of plant dispersal, competition and establishment, with an assumption of neutrality (all species had equivalent life histories) that reduced the initial dimensionality of the problem. The simulations assumed that the probability of immigration was a linear function of mainland abundance and distance to islands, similar to the equilibrium theory of island biogeography and the unified neutral theory of biodiversity. The rate of increase in species richness was not constant across island sizes, declining as island area became very large. This pattern was explained by the spatial dynamics of colonization and establishment, a non-random process that cannot be explained by passive sampling alone. Simulations showed that population establishment depended critically on rare long-distance dispersal events while population persistence was achieved by the formation of aggregated species distributions that developed through restricted dispersal and local competitive interactions. Nevertheless, species richness always declined to a single species in the absence of disturbances, while up to 40 species could persist to 10,000 years when spatially dependent mortality was added. Further explorations with spatially explicit models will be required to fully appreciate the consequence of land use change and altered disturbance regimes on patterns of species distribution and the maintenance of diversity.