Population synchrony and environmental variation: an experimental demonstration

Understanding the causes of population synchrony is an important issue for population management. Its study in field populations involves disentangling the effects of dispersal and correlated environmental noise. Here we report on an experimental investigation of the synchronizing effects of noise in closed laboratory populations of a soil mite, Sancassania berlesei . Mite life-histories are highly plastic with respect to resource availability (which is a function of food supply and population density). By varying the food supply we imposed environmental variation. We show that (a) population synchrony is a function of environmental synchrony, (b) perceived population synchrony depends on the life-history stage counted, and (c) average population synchrony tends to be lower than environmental synchrony: even when populations were supplied with food with a correlation of 1.0, the correlation between populations was 0.63 (bootstrapped 95%CI 0.54–0.71). This supports recent theoretical work suggesting that the Moran theorem (indicating that population synchrony equals environmental synchrony) generally overestimates the population synchrony of nonlinear systems.     

The role of large environmental noise in masting: General model and example from pistachio trees

Masting is synchronous, highly variable reproduction in a plant population, or synchronized boom-bust cycles of reproduction. These pulses of resources have cascading effects through ecosystems, and thus it is important to understand where they come from. How does masting happen and synchronize? In this paper, we suggest a mechanism for this. The mechanism is inspired by data from a pistachio orchard, which suggest that large environmental noise may play a crucial role in inducing masting in plant populations such as pistachio. We test this idea through development and analysis of a mathematical model of plant reproduction. We start with a very simple model, and generalize it based on the current models of plant reproduction and masting. Our results suggest that large environmental noise may indeed be a crucial part of the mechanism of masting in certain types of plant populations, including pistachio. This is a specific example of an important functional consequence of the interactions between stochasticity and nonlinearity.     

Synchrony of spatial populations: heterogeneous population dynamics and reddened environmental noise

Many species exhibit widespread spatial synchrony in population fluctuations. This pattern is of great ecological interest and can be a source of concern when a species is rare or endangered. Moran’s theorem suggests that if two (or more) populations sharing a common linear density-dependence in the renewal process are disturbed with correlated noise, they will become synchronized with correlation matching the noise correlation. In this report, correlation of nonidentical populations that are described by linear and stationary autoregressive processes is analyzed. We show that the expected spatial synchrony between two populations can be decomposed into two multiplicative components. One is the demographic component related to the values of the autoregressive coefficients and the noise color. The other is the spatial correlation of the environmental colored noise. The main results are consistent with the predictions of previous experiments and simulations, and the importance of this report is to provide theoretical support.     

Innovativeness, population size and cumulative cultural evolution

Henrich [Henrich, J., 2004. Demography and cultural evolution: how adaptive cultural processes can produce maladaptive losses—the Tasmanian case. Am. Antiquity 69, 197-214] proposed a model designed to show that larger population size facilitates cumulative cultural evolution toward higher skill levels. In this model, each newborn attempts to imitate the most highly skilled individual of the parental generation by directly-biased social learning, but the skill level he/she acquires deviates probabilistically from that of the exemplar (cultural parent). The probability that the skill level of the imitator exceeds that of the exemplar can be regarded as the innovation rate. After reformulating Henrich’s model rigorously, we introduce an overlapping-generations analog based on the Moran model and derive an approximate formula for the expected change per generation of the highest skill level in the population. For large population size, our overlapping-generations model predicts a much larger effect of population size than Henrich’s discrete-generations model. We then investigate by way of Monte Carlo simulations the case where each newborn chooses as his/her exemplar the most highly skilled individual from among a limited number of acquaintances. When the number of acquaintances is small relative to the population size, we find that a change in the innovation rate contributes more than a proportional change in population size to the cumulative cultural evolution of skill level.     

Dispersal, colored environmental noise, and spatial synchrony in population dynamics: analyzing a discrete host–parasitoid population model

Spatial synchrony is common, and its influences and causes have attracted the interest of ecologists. Spatially correlated environmental noise, dispersal, and trophic interactions have been considered as the causes of spatial synchrony. In this study, we developed a spatially structured population model, which is described by coupled-map lattices. Our recent investigation showed that trophic correlation of environmental noise was another important factor that affects spatial synchrony. As a supplement, we considered the influence of the color of the environmental noise on the spatial synchrony in this study. The noise color refers to the temporal correlation in the time series data of the noise, and is expressed as the degree of (first-order) autocorrelation for autoregressive noise. Patterns of spatial synchrony were considered for stable, periodic (quasi-periodic), and chaotic population dynamics. Numerical simulations verified that the color of the environmental noise is another mechanism that causes spatial synchrony. Generally, the effect of the color of the noise on the synchrony is dependent on the type of dynamics (stable, cyclic, chaotic) present in the population. For cyclic dynamics, simulation results clearly demonstrate that reddened noise has higher synchrony than white noise. The importance of our research is that it enriches the theory of potential causes of spatial synchrony.     

种群统计随机性和环境随机性对种群绝灭的影响

影响种群绝灭的随机干扰可分为种群统计随机性、环境随机性和随机灾害三大类。在相对稳定的环境条件下和相对较短的时间内,以前两类随机干扰对种群绝灭的影响为生态学家关注的焦点。但是,由于自然种群动态及其影响因子的复杂特征,进一步深入研究随机干扰对种群绝灭的作用在理论上和实践上都必须发展新的技术手段。本文回顾了种群统计随机性与环境随机性的概念起源与发展,系统阐述了其分析方法。归纳了两类随机性在种群绝灭研究中的应用范围、作用方式和特点的异同和区别方法。各类随机作用与种群动态之间关系的理论研究与对种群绝灭机理的实践研究紧密相关。根据理论模型模拟和自然种群实际分析两方面的研究现状,作者提出了进一步深入研究随机作用与种群非线性动态方法的策略。指出了随机干扰影响种群绝灭过程的研究的方向：更多的研究将从单纯的定性分析随机干扰对种群动力学简单性质的作用,转向结合特定的种群非线性动态特征和各类随机力作用特点具体分析绝灭极端动态的成因,以期做出精确的预测。     

Variation in dispersal mortality and dispersal propensity among individuals: the effects of age, sex and resource availability

1.  Dispersal of individuals between habitat patches depends on both the propensity to emigrate from a patch and the ability to survive inter-patch movement. Environmental factors and individual characteristics have been shown to influence dispersal rates but separating the effects of emigration and dispersal mortality on dispersal can often be difficult. In this study, we use a soil mite laboratory system to investigate factors affecting emigration and dispersal mortality.
2.  We tested the movement of different age groups in two-patch systems with different inter-patch distances. Differences in immigration among age groups were primarily driven by differences in emigration but dispersal mortality was greater for some groups. Immigration declined with increasing inter-patch distance, which was due to increasing dispersal mortality and decreasing emigration.
3.  In a second experiment, we compared the dispersal of recently matured males and females and tested the impact of food availability during the developmental period on their dispersal. Dispersal was found to be male biased but there was no significant sex bias in dispersal mortality. There was some evidence that food availability could affect emigration and dispersal mortality.
4.  These results demonstrate that both emigration and dispersal mortality can be affected by factors such as individual age and resource availability. Understanding these effects is likely to be important for predicting the fitness costs and population consequences of dispersal.     

MetaPopGen: an r package to simulate population genetics in large size metapopulations

Population genetics simulation models are useful tools to study the effects of demography and environmental factors on genetic variation and genetic differentiation. They allow for studying species and populations with complex life histories, spatial distribution and many other complicating factors that make analytical treatment impracticable. Most simulation models are individual‐based: this poses a limitation to simulation of very large populations because of the limits in computer memory and long computation times. To overcome these limitations, we propose an intermediate approach that allows modelling of very complex demographic scenarios, which would be intractable with analytical models, and removes the limitations imposed by large population size, which affect individual‐based simulation models. We implement this approach in a software package for the r environment, MetaPopGen. The innovative concept of this approach with respect to the other population genetic simulators is that it focuses on genotype numbers rather than on individuals. Genotype numbers are iterated through time by using random number generators for appropriate probabilistic distributions to reproduce the stochasticity inherent to Mendelian segregation, survival, dispersal and reproduction. Features included in the model are age structure, monoecious and dioecious (or separate sexes) life cycles, mutation, dispersal and selection. The model simulates only one locus at a time. All demographic parameters can be genotype‐, sex‐, age‐, deme‐ and time‐dependent. MetaPopGen is therefore indicated to study large populations and very complex demographic scenarios. We illustrate the capabilities of MetaPopGen by applying it to the case of a marine fish metapopulation in the Mediterranean Sea.     

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.     

Indirect evidence of density‐dependent population regulation in Aponomma hydrosauri (Acari: Ixodidae), an ectoparasite of reptiles

Abstract The extent to which density‐dependent processes regulate natural populations is the subject of an ongoing debate. We contribute evidence to this debate showing that density‐dependent processes influence the population dynamics of the ectoparasite Aponomma hydrosauri (Acari: Ixodidae), a tick species that infests reptiles in Australia. The first piece of evidence comes from an unusually long‐term dataset on the distribution of ticks among individual hosts. If density‐dependent processes are influencing either host mortality or vital rates of the parasite population, and those distributions can be approximated with negative binomial distributions, then general host–parasite models predict that the aggregation coefficient of the parasite distribution will increase with the average intensity of infections. We fit negative binomial distributions to the frequency distributions of ticks on hosts, and find that the estimated aggregation coefficient k increases with increasing average tick density. This pattern indirectly implies that one or more vital rates of the tick population must be changing with increasing tick density, because mortality rates of the tick's main host, the sleepy lizard, Tiliqua rugosa, are unaffected by changes in tick burdens. Our second piece of evidence is a re‐analysis of experimental data on the attachment success of individual ticks to lizard hosts using generalized linear modelling. The probability of successful engorgement decreases with increasing numbers of ticks attached to a host. This is direct evidence of a density‐dependent process that could lead to an increase in the aggregation coefficient of tick distributions described earlier. The population‐scale increase in the aggregation coefficient is indirect evidence of a density‐dependent process or processes sufficiently strong to produce a population‐wide pattern, and thus also likely to influence population regulation. The direct observation of a density‐dependent process is evidence of at least part of the responsible mechanism.     

The equilibrium population size of a partially migratory population and its response to environmental change

A partially migratory population consists of non‐migrant and migrant individuals that share a common site during one period of the annual cycle. In this paper, we derive the expected equilibrium population sizes of migrants and non‐migrants and show how the abundance of one type is dependent on the other because their dynamics are coupled through density‐dependent effects. We present an approach for developing hypotheses about how changes in the environment will influence partially migratory populations and for formulating testable predictions about the effects of future changes on the proportions of migrants and non‐migrants. We apply this approach to a hypothesis put forward by Berthold that improved environmental conditions at the shared site will generally increase the number of non‐migrants and decrease the number of migrants, and to a study by Nilsson et al. which observed an increase in the number of migrants in a partially migratory blue tit Cyanistes caeruleus population in Sweden, but an overall maintenance of the proportion of migrants.     

Demography_Lab,an educational application to evaluate population growth: Unstructured and matrix models

Training in Population Ecology asks for scalable applications capable of embarking students on a trip from basic concepts to the projection of populations under the various effects of density dependence and stochasticity. Demography_Lab is an educational tool for teaching Population Ecology aspiring to cover such a wide range of objectives. The application uses stochastic models to evaluate the future of populations. Demography_Lab may accommodate a wide range of life cycles and can construct models for populations with and without an age or stage structure. Difference equations are used for unstructured populations and matrix models for structured populations. Both types of models operate in discrete time. Models can be very simple, constructed with very limited demographic information or parameter‐rich, with a complex density‐dependence structure and detailed effects of the different sources of stochasticity. Demography_Lab allows for deterministic projections, asymptotic analysis, the extraction of confidence intervals for demographic parameters, and stochastic projections. Stochastic population growth is evaluated using up to three sources of stochasticity: environmental and demographic stochasticity and sampling error in obtaining the projection matrix. The user has full control on the effect of stochasticity on vital rates. The effect of the three sources of stochasticity may be evaluated independently for each vital rate. The user has also full control on density dependence. It may be included as a ceiling population size controlling the number of individuals in the population or it may be evaluated independently for each vital rate. Sensitivity analysis can be done for the asymptotic population growth rate or for the probability of extinction. Elasticity of the probability of extinction may be evaluated in response to changes in vital rates, and in response to changes in the intensity of density dependence and environmental stochasticity.     

Distribution of foundress group size in Ropalidia fasciata and R. plebeiana (Hymenoptera: Vespidae): an analysis using zero-truncated distribution models

Distribution patterns of the number of foundresses per newly established nest (foundress group size, FGS) of two primitively eusocial, independent-founding wasps, Ropalidia fasciata and R. plebeiana, were studied using zero-truncated distribution models. The distribution pattern of the FGS of R. fasciata is significantly different from a zero-truncated Poisson distribution but fits the zero-truncated negative binomial distribution well, indicating a strongly contagious distribution. R. plebeiana sometimes establishes new colonies by reusing old nests. In this case, distributions are strongly contagious. Competition among foundresses may be one reason for the contagious distribution of FGS in R. fasciata and in cases of old-nest reuse by R. plebeiana, but further studies, especially on the behaviour of foundresses in relation to FGS, are necessary. Electronic Publication     

How noise and coupling influence leading indicators of population extinction in a spatially extended ecological system

Anticipating critical transitions in spatially extended systems is a key topic of interest to ecologists. Gradually declining metapopulations are an important example of a spatially extended biological system that may exhibit a critical transition. Theory for spatially extended systems approaching extinction that accounts for environmental stochasticity and coupling is currently lacking. Here, we develop spatially implicit two-patch models with additive and multiplicative forms of environmental stochasticity that are slowly forced through population collapse, through changing environmental conditions. We derive patch-specific expressions for candidate indicators of extinction and test their performance via a simulation study. Coupling and spatial heterogeneities decrease the magnitude of the proposed indicators in coupled populations relative to isolated populations, and the noise regime and the degree of coupling together determine trends in summary statistics. This theory may be readily applied to other spatially extended ecological systems, such as coupled infectious disease systems on the verge of elimination.     

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

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Simultaneous confidence intervals for a success probability and intraclass correlation,with an application to screening mammography

This paper provides asymptotic simultaneous confidence intervals for a success probability and intraclass correlation of the beta‐binomial model, based on the maximum likelihood estimator approach. The coverage probabilities of those intervals are evaluated. An application to screening mammography is presented as an example. The individual and simultaneous confidence intervals for sensitivity and specificity and the corresponding intraclass correlations are investigated. Two additional examples using influenza data and sex ratio data among sibships are also considered, where the individual and simultaneous confidence intervals are provided.     

The amplification of environmental noise in population models: causes and consequences

Environmental variability is a ubiquitous feature of every organism's habitat. However, the interaction between density dependence and those density-independent factors that are manifested as environmental noise is poorly understood. We are interested in the conditions under which noise interacts with the density dependence to cause amplification of that noise when filtered by the system. For a broad family of structured population models, we show that amplification occurs near the threshold from stable to unstable dynamics by deriving an analytic formula for the amplification under weak noise. We confirm that the effect of noise is to sustain oscillations that would otherwise decay, and we show that it is the amplitude and not the phase that is affected. This is a feature noted in several recent studies. We study this phenomenon in detail for the lurchin and LPA models of population dynamics. We find that the degree of amplification is sensitive to both the noise input and life-history stage through which it acts, that the results hold for surprisingly high levels of noise, and that stochastic chaos (as measured by local Lyapunov exponents) is a concomitant feature of amplification. Further, it is shown that the temporal autocorrelation, or "color," of the noise has a major impact on the system response. We discuss the conditions under which color increases population variance and hence the risk of extinction, and we show that periodicity is sharpened when the color of the noise and dynamics coincide. Otherwise, there is interference, which shows how difficult it is in practice to separate the effects of nonlinearity and noise in short time series. The sensitivity of the population dynamics to noise when close to a bifurcation has wide-ranging consequences for the evolution and ecology of population dynamics.     

Evolution of variance in offspring number: The effects of population size and migration

It was shown by Gillespie [1974. Am. Nat. 108, 145–151], that if two genotypes produce the same average number of offspring on but have a different variance associated within each generation, the genotype with a lower variance will have a higher effective fitness. Specifically, the effective fitness is {ei65-1}, where w is the mean fitness, {ei65-2} is the variance in offspring number, and N is the total population size. The model also predicts that if a strategy has a higher arithmetic mean fitness and a higher variance than the competitor, the outcome of selection will depend on the population size (with larger population sizes favoring the highvariance, high-mean genotype). This suggests that for metapopulation sizes favoring the high-variance, high-mean genotype). This suggests that for metapopulations with large numbers of (relatively) small demes, a strategy with lower variance and lower mean may be favored if the migration rate is low while higher migration rates (consistent with a larger effective population size) favor the opposite strategy. Individual-based simulation confirms that this is indeed the case for an island model of migration, though the effect of migration differs greatly depending on whether migration precedes or follows selection. It is noted in the appendix that while Gillespie [1974. Am. Nat. 108, 145–151] does seem to be heuristically accurate, it is not clear that the definition of effective fitness follows from his derivation.