Spatial dynamics in the absence of dispersal: acorn production by oaks in central coastal California

We examined spatial patterns and spatial autocorrelation (synchrony) of annual acorn production in three species of oaks (genus Quercus ) over A 288 km transect in central coastal California. Over small (within-site) distances of <4 km, synchrony of acorn production between individual trees wits significant but varied through time and. for coast live oaks Q. agrifolia. differed al two sites 135 km apart. On a larger geographic scale, valley Q. lobata and blue Q. douglasii oaks exhibited significant synchrony in most distance categories between trees and sites up to 135 km apart and. in the case of coast live oaks, up to the maximum extent of the transect. Spatial patterns over this geographic scale also differed among species, with valley and blue oaks, but not coast live oaks, exhibiting distinct declines in synchrony of acorn production with distance. Interspecific synchrony in acorn production was generally lower than that within species but still significant over the entire extent of the survey. Spatial synchrony between sites was to some extent related to the same environmental variables previously found to correlate with annual acorn production within a site, suggesting that the environmental factors determining acorn production locally also influence spatial patterns over larger geographic areas. These results demonstrate that mast-fruiting in oaks occurs not only on a widespread geographic scale but also across species. They also confirm that synchrony over large geographic areas and complex spatial patterns varying in time can occur in systems where dispersal does not occur and thus environmental variability (the Moran effect) alone is likely to be driving spatial dynamics.     

Phase locking, the Moran effect and distance decay of synchrony: experimental tests in a model system

Spatially separated populations of many species fluctuate synchronously. Synchrony typically decays with increasing interpopulation distance. Spatial synchrony, and its distance decay, might reflect distance decay of environmental synchrony (the Moran effect), and/or short-distance dispersal. However, short-distance dispersal can synchronize entire metapopulations if within-patch dynamics are cyclic, a phenomenon known as phase locking. We manipulated the presence/absence of short-distance dispersal and spatially decaying environmental synchrony and examined their separate and interactive effects on the synchrony of the protist prey species Tetrahymena pyriformis growing in spatial arrays of patches (laboratory microcosms). The protist predator Euplotes patella consumed Tetrahymena and generated predator-prey cycles. Dispersal increased prey synchrony uniformly over both short and long distances, and did so by entraining the phases of the predator-prey cycles. The Moran effect also increased prey synchrony, but only over short distances where environmental synchrony was strongest, and did so by increasing the synchrony of stochastic fluctuations superimposed on the predator-prey cycle. Our results provide the first experimental demonstration of distance decay of synchrony due to distance decay of the Moran effect. Distance decay of the Moran effect likely explains distance decay of synchrony in many natural systems. Our results also provide an experimental demonstration of long-distance phase locking, and explain why cyclic populations provide many of the most dramatic examples of long-distance spatial synchrony in nature.     

Spatial synchrony in red grouse population dynamics

Red grouse Lagopus lagopus scoticus populations exhibit unstable dynamics that are often characterised by regular periodic fluctuations in abundance. Time-series' of grouse harvesting records collected from 287 management units (moors) across Scotland, England and Wales were analysed to investigate the broad scale patterns of synchrony in these fluctuations. Estimation of the spatial autocorrelation of grouse population dynamics across moors indicates relatively high levels of synchrony between populations on adjacent moors, but that this synchrony declines sharply with increasing inter-moor distance. At distances of greater than 100  km, grouse population time-series exhibit only weakly positive cross-correlation coefficients. Twenty-eight geographical, environmental and other candidate variables were examined to construct a general linear model to explain variation in local synchrony. Grouse moor productivity (average size of shooting bag), distance from the Atlantic coast moving in a north-easterly direction, April and June temperatures, and June rainfall significantly increased the explanatory power of this model. An understanding of the processes underlying synchrony in red grouse population dynamics is a prerequisite to anticipating the effects of large-scale environmental change on regional patterns of grouse distribution and abundance.     

The Moran effect revisited: spatial population synchrony under global warming

The world is spatially autocorrelated. Both abiotic and biotic properties are more similar among neighboring than distant locations, and their temporal co-fluctuations also decrease with distance. P. A. P. Moran realized the ecological importance of such ‘spatial synchrony’ when he predicted that isolated populations subject to identical log-linear density-dependent processes should have the same correlation in fluctuations of abundance as the correlation in environmental noise. The contribution from correlated weather to synchrony of populations has later been coined the ‘Moran effect’. Here, we investigate the potential role of the Moran effect in large-scale ecological outcomes of global warming. Although difficult to disentangle from dispersal and species interaction effects, there is compelling evidence from across taxa and ecosystems that spatial environmental synchrony causes population synchrony. Given this, and the accelerating number of studies reporting climate change effects on local population dynamics, surprisingly little attention has been paid to the implications of global warming for spatial population synchrony. However, a handful of studies of insects, birds, plants, mammals and marine plankton indicate decadal-scale changes in population synchrony due to trends in environmental synchrony. We combine a literature review with modeling to outline potential pathways for how global warming, through changes in the mean, variability and spatial autocorrelation of weather, can impact population synchrony over time. This is particularly likely under a ‘generalized Moran effect’, i.e. when relaxing Moran's strict assumption of identical log-linear density-dependence, which is highly unrealistic in the wild. Furthermore, climate change can influence spatial population synchrony indirectly, through its effects on dispersal and species interactions. Because changes in population synchrony may cascade through food-webs, we argue that the (generalized) Moran effect is key to understanding and predicting impacts of global warming on large-scale ecological dynamics, with implications for extinctions, conservation and management.     

Geographical variation in the spatial synchrony of a forest-defoliating insect: isolation of environmental and spatial drivers

Despite the pervasiveness of spatial synchrony of population fluctuations in virtually every taxon, it remains difficult to disentangle its underlying mechanisms, such as environmental perturbations and dispersal. We used multiple regression of distance matrices (MRMs) to statistically partition the importance of several factors potentially synchronizing the dynamics of the gypsy moth, an invasive species in North America, exhibiting outbreaks that are partially synchronized over long distances (approx. 900 km). The factors considered in the MRM were synchrony in weather conditions, spatial proximity and forest-type similarity. We found that the most likely driver of outbreak synchrony is synchronous precipitation. Proximity played no apparent role in influencing outbreak synchrony after accounting for precipitation, suggesting dispersal does not drive outbreak synchrony. Because a previous modelling study indicated weather might indirectly synchronize outbreaks through synchronization of oak masting and generalist predators that feed upon acorns, we also examined the influence of weather and proximity on synchrony of acorn production. As we found for outbreak synchrony, synchrony in oak masting increased with synchrony in precipitation, though it also increased with proximity. We conclude that precipitation could synchronize gypsy moth populations directly, as in a Moran effect, or indirectly, through effects on oak masting, generalist predators or diseases.     

有色环境噪音对空间异质种群动态同步性的影响

随着种群动态和空间结构研究兴趣的增加,激发了大量的有关空间同步性的理论和实验的研究工作。空间种群的同步波动现象在自然界广泛存在,它的影响和原因引起了很多生态学家的兴趣。Moran定理是一个非常重要的解释。但以往的研究大多假设环境变化为空间相关的白噪音。越来越多的研究表明很多环境变化的时间序列具有正的时间自相关性,也就是说用红噪音来描述更加合理。因此,推广经典的Moran效应来处理空间相关红噪音的情形很有必要。利用线性的二阶自回归过程的种群模型,推导了两种群空间同步性与种群动态异质性和环境变化的时间相关性(即环境噪音的颜色)之间的关系。深入分析了种群异质性和噪音颜色对空间同步性的影响。结果表明种群动态异质性不利于空间同步性,但详细的关系比较复杂。而红色噪音的同步能力体现在两方面:一方面,本身的相关性对同步性有贡献;另一方面,环境变化时间相关性可以通过改变种群密度依赖来影响同步性,但对同步性的影响并无一致性的结论,依赖于种群的平均动态等因素。这些结果对理解同步性的机理、利用同步机理来制定物种保护策略和害虫防治都有重要的意义。     

森林害虫发生的空间模式研究进展

森林昆虫种群表现出多样的时空模式,空间同步性是其中最常见的。回顾了森林昆虫空间同步性的特点、形成机制及研究方法方面的进展。森林害虫发生的同步性是广泛存在的,但不同昆虫种类的同步性大小不同。空间同步性常随距离的增大而下降,还与时间尺度有关。Moran效应和扩散是解释空间同步性的两种主要机制,通常Moran效应的影响要比扩散大。从虫害发生数据的获取、同步性的度量及成因3个方面介绍了空间同步性的研究方法方面的进展。利用树轮生态学原理重建森林虫害发生历史的方法可在事后获取可靠的数据,很值得国内研究者借鉴和应用。在空间自相关度量上,空间统计学方法和地统计学方法都是非常有力的手段,但由于不能处理多时间点数据而限制了其在同步性研究中的应用。在同步性成因研究中,利用变异分解将基于距离的Moran特征向量图(dbMEM)作为空间变量研究害虫发生的驱动力是比较新颖的研究方法。     

Generalizations of the Moran effect explaining spatial synchrony in population fluctuations

The Moran effect for populations separated in space states that the autocorrelations in the population fluctuations equal the autocorrelation in environmental noise, assuming the same linear density regulation in all populations. Here we generalize the Moran effect to include also nonlinear density regulation with spatial heterogeneity in local population dynamics as well as in the effects of environmental covariates by deriving a simple expression for the correlation between the sizes of two populations, using diffusion approximation to the theta-logistic model. In general, spatial variation in parameters describing the dynamics reduces population synchrony. We also show that the contribution of a covariate to spatial synchrony depends strongly on spatial heterogeneity in the covariate or in its effect on local dynamics. These analyses show exactly how spatial environmental covariation can synchronize fluctuations of spatially segregated populations with no interchange of individuals even if the dynamics are nonlinear.     

Geographically partitioned spatial synchrony among cyclic moth populations

Many species of forest lepidopterans exhibit regular population cycles, which culminate in outbreak densities at approximately ten-year intervals. Population peaks and mass outbreaks typically occur synchronously and may lead to extensive forest damages over large geographic areas. Here, we report patterns of spatial synchrony among cyclic autumnal moth ( Epirrita autumnata ) populations across Fennoscandia, as inferred from 24 long-term (10–33 years) data sets. The study provides the first formal analysis of spatial synchrony of this pest species which damages mountain birch ( Betula pubescens ssp. czerepanovii ) forests in the sub Arctic. We detected positive cross-correlations in population growth rates between the time series, indicating overall spatial synchrony. However, we found the strongest degree of synchrony within geographically and climatically distinct regional clusters, into which time series were partitioned using cluster analyses. Within regional clusters, moth populations were exposed to the synchronizing effects of common, spatially autocorrelated environmental conditions, i.e. a Moran effect. Consequently, we conclude that a geographically and climatically restricted Moran effect, perhaps interacting with dispersal, is the most likely explanation for the regionally partitioned pattern of synchrony among autumnal moth populations in Fennoscandia. Our results emphasize that high amounts of environmental variation may result in a clear structuring of spatial synchrony at unexpectedly small scales.     

A simultaneous test of synchrony causal factors in muskrat and mink fur returns at different scales across Canada

BACKGROUND: Synchrony among populations has been attributed to three major hypotheses: dispersal, the Moran effect, and trophic-level interactions. Unfortunately, simultaneous testing of these hypotheses demands complete and detailed data, which are scarce for ecological systems. METHODOLOGY/PRINCIPAL FINDINGS: Hudson's Bay Company data on mink and muskrat fur returns in Canada represent an excellent opportunity to test these hypotheses because of the detailed spatial and temporal data from this predator-prey system. Using structural equation modelling, support for each hypothesis was evaluated at two spatial scales: across Canada and dividing the country into three regions longitudinally. Our results showed that at both scales mink synchrony is a major factor determining muskrat synchrony, supporting the hypothesis of trophic-level interactions, but the influence of winter precipitation synchrony is also important in eastern Canada. Moreover, mink synchrony is influenced principally by winter precipitation synchrony at the level of all Canada (Moran effect), but by distance at regional level, which might suggest some influence of dispersal at this level. DISCUSSION/SIGNIFICANCE: Our result is one of the few reports of synchrony mediated by trophic-level interactions, highlighting the importance of evaluation of scale effects in population synchrony studies.     

Similarity in seasonal flow regimes,not regional environmental classifications explain synchrony in brown trout population dynamics in France

相似文献   

Environmental colour intensifies the Moran effect when population dynamics are spatially heterogeneous

Evidence for synchronous fluctuations of spatially separated populations is ubiquitous in the literature, including accounts within and across taxa. Among the few mechanisms explaining this phenomenon is the Moran effect, whereby independent populations are synchronized by spatially correlated environmental disturbances. The body of research on the Moran effect predominantly assumes that environmental disturbances within a local site are serially uncorrelated; that is, successive observations in time at a particular local site are independent. Yet, many environmental variables are known to possess strong temporal autocorrelation – a character which has often been described as 'colour'. The omission of environmental colour from research on the Moran effect may be due in part to the lack of methods capable of generating sets of time series with a desired colour and spatial correlation. Here I present a novel and simple method designated as 'phase partnering' to generate such sets of time series and I investigate the combined impact of spatial correlation and environmental colour on population synchrony in two common models of population dynamics. For linear population dynamics, and for a subset of nonlinear population dynamics, coloured environments intensify the Moran effect when population dynamics are spatially heterogeneous; in coloured environments the spatial correlation between populations more closely mimics the spatial correlation between their respective environments. Given that most environmental variables are coloured, these results imply that the Moran effect may be a far more significant driver of regional-scale population and interspecific synchrony than is currently believed.     

Does the pattern of population synchrony through space reveal if the Moran effect is acting?

The populations of many species fluctuate in synchrony across large geographical areas. This synchrony is often attributed to the Moran effect, that is, shared environmental fluctuations across the region. In this article, I use a series of simple metapopulation models to show that the degree of synchrony among populations separated by different distances is strongly affected by the particular way that environmental stochasticity is represented in the models. Furthermore, when multiple types of stochasticity are acting simultaneously, the synchronizing effect of any one type is difficult to discern from the resulting pattern of population synchrony. These effects can be exacerbated under certain demographic conditions or if population dynamics are affected by interspecific interactions. In general, it should be extremely difficult to determine if synchrony is caused by the Moran effect using only the synchrony–distance relationship of natural populations.     

Spatial heterogeneity in climate change effects decouples the long‐term dynamics of wild reindeer populations in the high Arctic

The ‘Moran effect’ predicts that dynamics of populations of a species are synchronized over similar distances as their environmental drivers. Strong population synchrony reduces species viability, but spatial heterogeneity in density dependence, the environment, or its ecological responses may decouple dynamics in space, preventing extinctions. How such heterogeneity buffers impacts of global change on large‐scale population dynamics is not well studied. Here, we show that spatially autocorrelated fluctuations in annual winter weather synchronize wild reindeer dynamics across high‐Arctic Svalbard, while, paradoxically, spatial variation in winter climate trends contribute to diverging local population trajectories. Warmer summers have improved the carrying capacity and apparently led to increased total reindeer abundance. However, fluctuations in population size seem mainly driven by negative effects of stochastic winter rain‐on‐snow (ROS) events causing icing, with strongest effects at high densities. Count data for 10 reindeer populations 8–324 km apart suggested that density‐dependent ROS effects contributed to synchrony in population dynamics, mainly through spatially autocorrelated mortality. By comparing one coastal and one ‘continental’ reindeer population over four decades, we show that locally contrasting abundance trends can arise from spatial differences in climate change and responses to weather. The coastal population experienced a larger increase in ROS, and a stronger density‐dependent ROS effect on population growth rates, than the continental population. In contrast, the latter experienced stronger summer warming and showed the strongest positive response to summer temperatures. Accordingly, contrasting net effects of a recent climate regime shift—with increased ROS and harsher winters, yet higher summer temperatures and improved carrying capacity—led to negative and positive abundance trends in the coastal and continental population respectively. Thus, synchronized population fluctuations by climatic drivers can be buffered by spatial heterogeneity in the same drivers, as well as in the ecological responses, averaging out climate change effects at larger spatial scales.     

The extended Moran effect and large-scale synchronous fluctuations in the size of great tit and blue tit populations

1. Synchronous fluctuations of geographically separated populations are in general explained by the Moran effect, i.e. a common influence on the local population dynamics of environmental variables that are correlated in space. Empirical support for such a Moran effect has been difficult to provide, mainly due to problems separating out effects of local population dynamics, demographic stochasticity and dispersal that also influence the spatial scaling of population processes. Here we generalize the Moran effect by decomposing the spatial autocorrelation function for fluctuations in the size of great tit Parus major and blue tit Cyanistes caeruleus populations into components due to spatial correlations in the environmental noise, local differences in the strength of density regulation and the effects of demographic stochasticity. 2. Differences between localities in the strength of density dependence and nonlinearity in the density regulation had a small effect on population synchrony, whereas demographic stochasticity reduced the effects of the spatial correlation in environmental noise on the spatial correlations in population size by 21.7% and 23.3% in the great tit and blue tit, respectively. 3. Different environmental variables, such as beech mast and climate, induce a common environmental forcing on the dynamics of central European great and blue tit populations. This generates synchronous fluctuations in the size of populations located several hundred kilometres apart. 4. Although these environmental variables were autocorrelated over large areas, their contribution to the spatial synchrony in the population fluctuations differed, dependent on the spatial scaling of their effects on the local population dynamics. We also demonstrate that this effect can lead to the paradoxical result that a common environmental variable can induce spatial desynchronization of the population fluctuations. 5. This demonstrates that a proper understanding of the ecological consequences of environmental changes, especially those that occur simultaneously over large areas, will require information about the spatial scaling of their effects on local population dynamics.     

Life history strategies affect climate based spatial synchrony in population dynamics of West African freshwater fishes

Spatial synchrony in species abundance is a general phenomenon that has been found in populations representing virtually all major taxa. Dispersal among populations and synchronous stochastic effects (the so called "Moran effect") are the mechanisms most likely to explain such synchrony patterns. Very few studies have related the degree of spatial synchrony to the biological characteristics of species. Here we present a case where specific predictions can be made to relate river fish species characteristics and synchrony determined exclusively by a Moran effect through the expected sensitivity of species to the regional component of environmental stochasticity. By analyzing 23-year time series of abundance estimates in two isolated localities we show that species associated with synchronized reproduction during the wet season, high fecundity, small egg size and high gonado-somatic index (the so called "periodic" strategy) have a higher degree of spatial synchrony in population dynamics than species associated with the opposite traits (the so called "equilibrium" strategy). This is supported by significant relationships (P values <0.01) between species traits and the levels of synchrony after removing taxonomical relatedness. Spatial synchrony computed from summed annual total catches by groups of species, separated into strategy types also showed a significantly higher degree of synchrony for the periodic (r=0.83) than the equilibrium (r=0.46) group. Regional hydrological variability is likely to be partly responsible for the observed synchrony pattern and a regional discharge index showed better relationships with the periodic group, supporting the expected differential effect of regional environmental correlation on population dynamics.     

Travelling wave dynamics and self-organization in a spatio-temporally structured population

We analyse spatial population dynamics showing that periodic or period-like chaotic dynamics produce self-organization structures, such as travelling waves. We suggest that self-organized patterns are associated with spatial synchrony patterns that often depend on geographical distance between subpopulations. The population dynamics also show statistical spatial autocorrelation patterns. We contrast our theoretical simulations with empirical data on annual damages in young sapling stands caused by voles. We conclude, on the basis of the periodicity, synchrony, and spatial autocorrelation patterns, and our simulation results, that vole dynamics represent travelling waves in population dynamics. We suggest that because such synchrony patterns are frequently observed in natural populations, spatial self-organization may be more common in population dynamics than reported in the literature.     

Testing Moran's theorem in an agroecosystem

The abundance and reproductive effort of populations frequently fluctuate across space and time, a phenomenon known as spatial synchrony. Knowledge of the causes of this behavior underlies the ability to manage species, protect the health of humans and the environment, and increase agricultural sustainability. We used an agroecosystem to test Moran's theorem – spatial synchrony results from environmental entrainment. The controlled conditions of the agroecosystem allowed us to create a highly correlated environment while negating the effects of the alternative hypotheses: dispersal and trophic interactions. Under such conditions, synchrony of fruit production by 4288 trees was high over six years in a 32.5 ha pistachio orchard and occurred at similar temporal frequency as weather patterns demonstrating the Moran effect. The spatial synchrony of fruit production was less than the presumed synchrony of the environment supporting research from microcosms and observational studies showing the Moran effect is degraded by local mechanisms. Indeed even under the homogeneous environment of this system, synchrony declined significantly with distance among trees. We present evidence suggesting that the correlation of the local environment affects intrinsic dynamics to cause these patterns. Our findings demonstrate that the Moran effect is, at minimum, partially responsible for the synchronous fruit production in this system. Agroecosystems are often overlooked in basic ecological research; this experiment provides an example of their comparative advantages for the study of some ecological questions.     

Patterns of Annual Seed Production by Northern Hemisphere Trees: A Global Perspective

We tested whether annual seed production (masting or mast fruiting) in Northern Hemisphere trees is an evolved strategy or a consequence of resource tracking by comparing masting patterns with those of annual rainfall and mean summer temperatures, two environmental variables likely to correlate with available resources. There were generally significant negative autocorrelations between the seed crop in year x and year x+1 (year x+2 in species of Quercus requiring 2 yr to mature acorns), as expected if resources are depleted in mast years in part by switching resources from growth to reproduction. Spatial autocorrelation in annual seed production generally declined with distance but was statistically significant over large geographic areas. Variability in annual seed production was relatively high and inversely correlated with latitude and generally not bimodally distributed. Patterns of spatial autocorrelation in annual rainfall and summer temperatures are generally similar to those exhibited by annual seed production, and relative variability in annual rainfall is also inversely correlated with latitude. However, these environmental variables exhibit distinctly different patterns of temporal autocorrelation, are much less variable, and are more normally distributed than annual seed production. Combined with the inverse relationship between growth and reproduction previously documented, these results support the hypothesis that variability in annual seed production is an evolved strategy and that annual seed production is more or less normally distributed rather than an all-or-none phenomenon.     

Spatial synchrony in population fluctuations: extending the Moran theorem to cope with spatially heterogeneous dynamics

The recent interest in the spatial structure and dynamics of populations motivated numerous theoretical and empirical studies of spatial synchrony, the tendency of populations to fluctuate in unison over regional areas. The first comprehensive framework applied to spatial synchrony was probably the one elaborated by P. A. P. Moran back in 1953. He suggested that if two populations have the same linear density-dependent structure, the correlation between them will be equal to that between the local density-independent conditions. Surprisingly, the consequences of violating the assumption that the dynamics of the populations are identical has received little attention. In this paper, making the assumption that population dynamics can be described by linear and stationary autoregressive processes, I show that the observed spatial synchrony between two populations can be decomposed into two multiplicative components: the demographic component depending on the values of the autoregressive coefficients, and the correlation of the environmental noise. The Moran theorem corresponds to the special case where the demographic component equals unity. Using published data, I show that the spatial variability in population dynamics may substantially contribute to the spatial variability of population synchrony, and thus should not be neglected in future studies.