Rapid surrogate testing of wavelet coherences

Background. The use of wavelet coherence methods enables the identification of frequency-dependent relationships between the phases of the fluctuations found in complex systems such as medical and other biological timeseries. These relationships may illuminate the causal mechanisms that relate the variables under investigation. However, computationally intensive statistical testing is required to ensure that apparent phase relationships are statistically significant, taking into account the tendency for spurious phase relationships to manifest in short stretches of data. Methods. In this study we revisit Fourier transform based methods for generating surrogate data, with which we sample the distribution of coherence values associated with the null hypothesis that no actual phase relationship between the variables exists. The properties of this distribution depend on the cross-spectrum of the data. By describing the dependency, we demonstrate how large numbers of values from this distribution can be rapidly generated without the need to generate correspondingly many wavelet transforms. Results. As a demonstration of the technique, we apply the efficient testing methodology to a complex biological system consisting of population timeseries for planktonic organisms in a food web, and certain environmental drivers. A large number of frequency dependent phase relationships are found between these variables, and our algorithm efficiently determines the probability of each arising under the null hypothesis, given the length and properties of the data. Conclusion. Proper accounting of how bias and wavelet coherence values arise from cross spectral properties provides a better understanding of the expected results under the null hypothesis. Our new technique enables enormously faster significance testing of wavelet coherence

Climate-change related regime shifts have altered spatial synchrony of plankton dynamics in the North Sea

During the 1980s the North Sea plankton community underwent a well-documented ecosystem regime shift, including both spatial changes (northward species range shifts) and temporal changes (increases in the total abundances of warmer-water species). This regime shift has been attributed to climate change. Plankton provide a link between climate and higher-trophic-level organisms, which can forage on large spatial and temporal scales. It is therefore important to understand not only whether climate change affects purely spatial or temporal aspects of plankton dynamics, but also whether it affects spatio-temporal aspects such as metapopulation synchrony. If plankton synchrony is altered, higher-trophic-level feeding patterns may be modified. A second motivation for investigating changes in synchrony is that the possibility of such alterations has been examined for few organisms, in spite of the fact that synchrony is ubiquitous and of major importance in ecology. This study uses correlation coefficients and spectral analysis to investigate whether synchrony changed between the periods 1959-1980 and 1989-2010. Twenty-three plankton taxa, sea surface temperature (SST), and wind speed were examined. Results revealed that synchrony in SST and plankton was altered. Changes were idiosyncratic, and were not explained by changes in abundance. Changes in the synchrony of Calanus helgolandicus and Para-pseudocalanus spp appeared to be driven by changes in SST synchrony. This study is one of few to document alterations of synchrony and climate-change impacts on synchrony. We discuss why climate-change impacts on synchrony may well be more common and consequential than previously recognized

Synchrony affects Taylor’s law in theory and data

Two widely confirmed patterns in ecology are Taylor’s law (TL), which states that the variance of population density is approximately a power of mean population density, and population synchrony, the tendency of species’ population sizes in different areas to be correlated through time. TL has been applied in many areas, including fisheries management, conservation, agriculture, finance, physics, and meteorology. Synchrony of populations increases the likelihood of large-scale pest or disease outbreaks and shortages of resources. We show that changed synchrony modifies and can invalidate TL. Widespread recent changes in synchrony, possibly resulting from climate change, may broadly affect TL and its applications

Synchrony is more than its top-down and climatic parts: interacting Moran effects on phytoplankton in British seas

Large-scale spatial synchrony is ubiquitous in ecology. We examined 56 years of data representing chlorophyll density in 26 areas in British seas monitored by the Continuous Plankton Recorder survey. We used wavelet methods to disaggregate synchronous fluctuations by timescale and determine that drivers of synchrony include both biotic and abiotic variables. We tested these drivers for statistical significance by comparison with spatially synchronous surrogate data. We generated timescale-specific models, accounting for 61% of long-timescale (> 4yrs) synchrony in a chlorophyll density index, but only 3% of observed short-timescale (< 4yrs) synchrony. The dominant source of long-timescale chlorophyll synchrony was closely related to sea surface temperature, through a Moran effect, though likely via complex oceanographic mechanisms. The top-down action of Calanus finmarchicus predation enhances this environmental synchronising mechanism and interacts with it non-additively to produce more long-timescale synchrony than top-down and climatic drivers would produce independently. Thus we demonstrate interaction effects between Moran drivers of synchrony, a new mechanism for synchrony that may affect many ecosystems at large spatial scales

A new approach to interspecific synchrony in population ecology using tail association

AbstractStandard methods for studying the association between two ecologically important variables provide only a small slice of the information content of the association, but statistical approaches are available that provide comprehensive information. In particular, available approaches can reveal tail associations, that is, accentuated or reduced associations between the more extreme values of variables. We here study the nature and causes of tail associations between phenological or population‐density variables of co‐located species, and their ecological importance. We employ a simple method of measuring tail associations which we call the partial Spearman correlation. Using multidecadal, multi‐species spatiotemporal datasets on aphid first flights and marine phytoplankton population densities, we assess the potential for tail association to illuminate two major topics of study in community ecology: the stability or instability of aggregate community measures such as total community biomass and its relationship with the synchronous or compensatory dynamics of the community's constituent species; and the potential for fluctuations and trends in species phenology to result in trophic mismatches. We find that positively associated fluctuations in the population densities of co‐located species commonly show asymmetric tail associations; that is, it is common for two species’ densities to be more correlated when large than when small, or vice versa. Ordinary measures of association such as correlation do not take this asymmetry into account. Likewise, positively associated fluctuations in the phenology of co‐located species also commonly show asymmetric tail associations. We provide evidence that tail associations between two or more species’ population‐density or phenology time series can be inherited from mutual tail associations of these quantities with an environmental driver. We argue that our understanding of community dynamics and stability, and of phenologies of interacting species, can be meaningfully improved in future work by taking into account tail associations.</jats:p

How environmental drivers of spatial synchrony interact

Spatial synchrony, the tendency for populations across space to show correlated fluctuations, is a fundamental feature of population dynamics, linked to central topics of ecology such as population cycling, extinction risk, and ecosystem stability. A common mechanism of spatial synchrony is the Moran effect, whereby spatially synchronized environmental signals drive population dynamics and hence induce population synchrony. After reviewing recent progress in understanding Moran effects, we here elaborate a general theory of how Moran effects of different environmental drivers acting on the same populations can interact, either synergistically or destructively, to produce either substantially more or markedly less population synchrony than would otherwise occur. We provide intuition for how this newly recognized mechanism works through theoretical case studies and application of our theory to California populations of giant kelp. We argue that Moran interactions should be common. Our theory and analysis explain an important new aspect of a fundamental feature of spatiotemporal population dynamics

Tail dependent spatial synchrony arises from nonlinear driver response relationships

Spatial synchrony may be tail-dependent, that is, stronger when populations are abundant than scarce, or vice-versa. Here, ‘tail-dependent’ follows from distributions having a lower tail consisting of relatively low values and an upper tail of relatively high values. We present a general theory of how the distribution and correlation structure of an environmental driver translates into tail-dependent spatial synchrony through a non-linear response, and examine empirical evidence for theoretical predictions in giant kelp along the California coastline. In sheltered areas, kelp declines synchronously (lower-tail dependence) when waves are relatively intense, because waves below a certain height do little damage to kelp. Conversely, in exposed areas, kelp is synchronised primarily by periods of calmness that cause shared recovery (upper-tail dependence). We find evidence for geographies of tail dependence in synchrony, which helps structure regional population resilience: areas where population declines are asynchronous may be more resilient to disturbance because remnant populations facilitate reestablishment

Moth biomass increases and decreases over 50 years in Britain

Steep insect biomass declines ('insectageddon') have been widely reported, despite a lack of continuously collected biomass data from replicated long-term monitoring sites. Such severe declines are not supported by the world’s longest running insect population database: annual moth biomass estimates from British fixed monitoring sites revealed increasing biomass between 1967 and 1982, followed by gradual decline from 1982 to 2017, with a 2.2-fold net gain in mean biomass between the first (1967–1976) and last decades (2008–2017) of monitoring. High between-year variability and multi-year periodicity in biomass emphasize the need for long-term data to detect trends and identify their causes robustly

An Emerging Infectious Disease Triggering Large-Scale Hyperpredation

Hyperpredation refers to an enhanced predation pressure on a secondary prey due to either an increase in the abundance of a predator population or a sudden drop in the abundance of the main prey. This scarcely documented mechanism has been previously studied in scenarios in which the introduction of a feral prey caused overexploitation of native prey. Here we provide evidence of a previously unreported link between Emergent Infectious Diseases (EIDs) and hyperpredation on a predator-prey community. We show how a viral outbreak caused the population collapse of a host prey at a large spatial scale, which subsequently promoted higher-than-normal predation intensity on a second prey from shared predators. Thus, the disease left a population dynamic fingerprint both in the primary host prey, through direct mortality from the disease, and indirectly in the secondary prey, through hyperpredation. This resulted in synchronized prey population dynamics at a large spatio-temporal scale. We therefore provide evidence for a novel mechanism by which EIDs can disrupt a predator-prey interaction from the individual behavior to the population dynamics. This mechanism can pose a further threat to biodiversity through the human-aided disruption of ecological interactions at large spatial and temporal scales.MM and JASZ were partially supported by a project of the Spanish Ministerio de Educación y Ciencia (reference CGL-2006-10689/BOS)

The long and the short of it: Mechanisms of synchronous and compensatory dynamics across temporal scales

Synchronous dynamics (fluctuations that occur in unison) are universal phenomena with widespread implications for ecological stability. Synchronous dynamics can amplify the destabilizing effect of environmental variability on ecosystem functions such as productivity, whereas the inverse, compensatory dynamics, can stabilize function. Here we combine simulation and empirical analyses to elucidate mechanisms that underlie patterns of synchronous versus compensatory dynamics. In both simulated and empirical communities, we show that synchronous and compensatory dynamics are not mutually exclusive but instead can vary by timescale. Our simulations identify multiple mechanisms that can generate timescale-specific patterns, including different environmental drivers, diverse life histories, dispersal, and non-stationary dynamics. We find that traditional metrics for quantifying synchronous dynamics are often biased toward long-term drivers and may miss the importance of short-term drivers. Our findings indicate key mechanisms to consider when assessing synchronous versus compensatory dynamics and our approach provides a pathway for disentangling these dynamics in natural systems