Opposing Patterns of Spatial Synchrony in Lyme Disease Incidence.

Incidence of Lyme disease, a tick-borne illness prevalent in the US, is increasing in endemic regions and regions with no previous history of the disease, significantly impacting public health. We examined space-time patterns of Lyme disease incidence and the influence of ecological and social factors on spatial synchrony, i.e., correlated incidence fluctuations across US counties. Specifically, we addressed these questions: Does Lyme disease incidence exhibit spatial synchrony? If so, what geographic patterns does Lyme disease synchrony exhibit? Are geographic patterns of disease synchrony related to weather, land cover, access to health care, or tick-borne disease awareness? How do effects of these variables on Lyme disease synchrony differ geographically? We used network analysis and matrix regression to examine geographical patterns of Lyme disease synchrony and their potential mechanisms in 399 counties in the eastern and Midwestern US. We found two distinct regions of synchrony in Northeast and upper Midwest regions exhibiting opposing temporal fluctuations in incidence. Spatial patterns of Lyme disease synchrony were partly explained by land cover, weather, poverty, and awareness of tick-borne illness, with significant predictive variables changing regionally. However, the two regions may have become more synchronous over time, potentially leading to higher-amplitude nation-wide fluctuations in disease incidence.

Geographic variation in larval cold tolerance and exposure across the invasion front of a widely established forest insect.

Under global climate change, high and low temperature extremes can drive shifts in species distributions. Across the range of a species, thermal tolerance is based on acclimatization, plasticity, and may undergo selection, shaping resilience to temperature stress. In this study, we measured variation in cold temperature tolerance of early instar larvae of an invasive forest insect, Lymantria dispar dispar L. (Lepidoptera: Erebidae), using populations sourced from a range of climates within the current introduced range in the Eastern United States. We tested for population differences in chill coma recovery (CCR) by measuring recovery time following a period of exposure to a nonlethal cold temperature in 2 cold exposure experiments. A 3rd experiment quantified growth responses after CCR to evaluate sublethal effects. Our results indicate that cold tolerance is linked to regional climate, with individuals from populations sourced from colder climates recovering faster from chill coma. While this geographic gradient is seen in many species, detecting this pattern is notable for an introduced species founded from a single point-source introduction. We demonstrate that the cold temperatures used in our experiments occur in nature during cold spells after spring egg hatch, but impacts to growth and survival appear low. We expect that population differences in cold temperature performance manifest more from differences in temperature-dependent growth than acute exposure. Evaluating intraspecific variation in cold tolerance increases our understanding of the role of climatic gradients on the physiology of an invasive species, and contributes to tools for predicting further expansion.

Dispersal synchronizes giant kelp forests.

Spatial synchrony is the tendency for population fluctuations to be correlated among different locations. This phenomenon is a ubiquitous feature of population dynamics and is important for ecosystem stability, but several aspects of synchrony remain unresolved. In particular, the extent to which any particular mechanism, such as dispersal, contributes to observed synchrony in natural populations has been difficult to determine. To address this gap, we leveraged recent methodological improvements to determine how dispersal structures synchrony in giant kelp (Macrocystis pyrifera), a global marine foundation species that has served as a useful system for understanding synchrony. We quantified population synchrony and fecundity with satellite imagery across 11 years and 880 km of coastline in southern California, USA, and estimated propagule dispersal probabilities using a high-resolution ocean circulation model. Using matrix regression models that control for the influence of geographic distance, resources (seawater nitrate), and disturbance (destructive waves), we discovered that dispersal was an important driver of synchrony. Our findings were robust to assumptions about propagule mortality during dispersal and consistent between two metrics of dispersal: (1) the individual probability of dispersal and (2) estimates of demographic connectivity that incorporate fecundity (the number of propagules dispersing). We also found that dispersal and environmental conditions resulted in geographic clusters with distinct patterns of synchrony. This study is among the few to statistically associate synchrony with dispersal in a natural population and the first to do so in a marine organism. The synchronizing effects of dispersal and environmental conditions on foundation species, such as giant kelp, likely have cascading effects on the spatial stability of biodiversity and ecosystem function.

Spatial synchrony cascades across ecosystem boundaries and up food webs via resource subsidies.

Cross-ecosystem subsidies are critical to ecosystem structure and function, especially in recipient ecosystems where they are the primary source of organic matter to the food web. Subsidies are indicative of processes connecting ecosystems and can couple ecological dynamics across system boundaries. However, the degree to which such flows can induce cross-ecosystem cascades of spatial synchrony, the tendency for system fluctuations to be correlated across locations, is not well understood. Synchrony has destabilizing effects on ecosystems, adding to the importance of understanding spatiotemporal patterns of synchrony transmission. In order to understand whether and how spatial synchrony cascades across the marine-terrestrial boundary via resource subsidies, we studied the relationship between giant kelp forests on rocky nearshore reefs and sandy beach ecosystems that receive resource subsidies in the form of kelp wrack (detritus). We found that synchrony cascades from rocky reefs to sandy beaches, with spatiotemporal patterns mediated by fluctuations in live kelp biomass, wave action, and beach width. Moreover, wrack deposition synchronized local abundances of shorebirds that move among beaches seeking to forage on wrack-associated invertebrates, demonstrating that synchrony due to subsidies propagates across trophic levels in the recipient ecosystem. Synchronizing resource subsidies likely play an underappreciated role in the spatiotemporal structure, functioning, and stability of ecosystems.

Climatic effects on the synchrony and stability of temperate headwater invertebrates over four decades.

Important clues about the ecological effects of climate change can arise from understanding the influence of other Earth-system processes on ecosystem dynamics but few studies span the inter-decadal timescales required. We, therefore, examined how variation in annual weather patterns associated with the North Atlantic Oscillation (NAO) over four decades was linked to synchrony and stability in a metacommunity of stream invertebrates across multiple, contrasting headwaters in central Wales (UK). Prolonged warmer and wetter conditions during positive NAO winters appeared to synchronize variations in population and community composition among and within streams thereby reducing stability across levels of ecological organization. This climatically mediated synchronization occurred in all streams irrespective of acid-base status and land use, but was weaker where invertebrate communities were more functionally diverse. Wavelet linear models indicated that variation in the NAO explained up to 50% of overall synchrony in species abundances at a timescale of 4-6 years. The NAO appeared to affect ecological dynamics through local variations in temperature, precipitation and discharge, but increasing hydrochemical variability within sites during wetter winters might have contributed. Our findings illustrate how large-scale climatic fluctuations generated over the North Atlantic can affect population persistence and dynamics in inland freshwater ecosystems in ways that transcend local catchment character. Protecting and restoring functional diversity in stream communities might increase their stability against warmer, wetter conditions that are analogues of ongoing climate change. Catchment management could also dampen impacts and provide options for climate change adaptation.

Seasonality in Environment and Population Processes Alters Population Spatial Synchrony.

AbstractPopulation spatial synchrony-the tendency for temporal population fluctuations to be correlated across locations-is common and important to metapopulation stability and persistence. One common cause of spatial synchrony, termed the Moran effect, occurs when populations respond to environmental fluctuations, such as weather, that are correlated over space. Although the degree of spatial synchrony in environmental fluctuations can differ between seasons and different population processes occur in different seasons, the impact on population spatial synchrony is uncertain because prior work has largely assumed that the spatial synchrony of environmental fluctuations and their effect on populations are consistent over annual sampling intervals. We used theoretical models to examine how seasonality in population processes and the spatial synchrony of environmental drivers affect population spatial synchrony. We found that population spatial synchrony can depend not only on the spatial synchrony of environmental drivers but also on the degree to which environmental fluctuations are correlated across seasons, locally, and across space. Moreover, measurements of synchrony from "snapshot" population censuses may not accurately reflect synchrony during other parts of the year. Together, these results show that neglecting seasonality in environmental conditions and population processes is consequential for understanding population spatial synchrony and its driving mechanisms.

Advances in understanding the drivers of population spatial synchrony.

The causes of spatial synchrony in population dynamics are often elusive. We review how recent advances have enhanced understanding of the causes of the spatial synchrony of insect populations and revealed previously underappreciated complexities in patterns of synchrony. We highlight how regional-scale studies of population genetic structure have helped elucidate the role of dispersal in population synchronization and how novel data-analytic approaches have revealed variation in spatial synchrony across timescales and geographies and the underlying drivers. We also stress the limited current understanding of the impacts of climate change on the spatial synchrony of insect populations and the potential ramifications of these effects for pest management as well as species conservation.

Growth and development of an invasive forest insect under current and future projected temperature regimes.

Temperature and its impact on fitness are fundamental for understanding range shifts and population dynamics under climate change. Geographic climate heterogeneity, behavioral and physiological plasticity, and thermal adaptation to local climates make predicting the responses of species to climate change complex. Using larvae from seven geographically distinct wild populations in the eastern United States of the non-native forest pest Lymantria dispar dispar (L.), we conducted a simulated reciprocal transplant experiment in environmental chambers using six custom temperature regimes representing contemporary conditions near the southern and northern extremes of the US invasion front and projections under two climate change scenarios for the year 2050. Larval growth and development rates increased with climate warming compared with current thermal regimes and tended to be greater for individuals originally sourced from southern rather than northern populations. Although increases in growth and development rates with warming varied somewhat by region of the source population, there was not strong evidence of local adaptation, southern populations tended to outperform those from northern populations in all thermal regimes. Our study demonstrates the utility of simulating thermal regimes under climate change in environmental chambers and emphasizes how the impacts from future increases in temperature can vary based on geographic differences in climate-related performance among populations.

Disturbance and nutrients synchronise kelp forests across scales through interacting Moran effects.

Spatial synchrony is a ubiquitous and important feature of population dynamics, but many aspects of this phenomenon are not well understood. In particular, it is largely unknown how multiple environmental drivers interact to determine synchrony via Moran effects, and how these impacts vary across spatial and temporal scales. Using new wavelet statistical techniques, we characterised synchrony in populations of giant kelp Macrocystis pyrifera, a widely distributed marine foundation species, and related synchrony to variation in oceanographic conditions across 33 years (1987-2019) and >900 km of coastline in California, USA. We discovered that disturbance (storm-driven waves) and resources (seawater nutrients)-underpinned by climatic variability-act individually and interactively to produce synchrony in giant kelp across geography and timescales. Our findings demonstrate that understanding and predicting synchrony, and thus the regional stability of populations, relies on resolving the synergistic and antagonistic Moran effects of multiple environmental drivers acting on different timescales.

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.

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.

Assessing drivers of localized invasive spread to inform large-scale management of a highly damaging insect pest.

Studies of biological invasions at the macroscale or across multiple scales can provide important insights for management, particularly when localized information about invasion dynamics or environmental contexts is unavailable. In this study, we performed a macroscale analysis of the roles of invasion drivers on the local scale dynamics of a high-profile pest, Lymantria dispar dispar L., with the purpose of improving the prioritization of vulnerable areas for treatment. Specifically, we assessed the relative effects of various anthropogenic and environmental variables on the establishment rate of 8010 quadrats at a localized scale (5 × 5 km) across the entire L. dispar transition zone (the area encompassing the leading population edge, currently from Minnesota to North Carolina). We calculated the number of years from first detection of L. dispar in a quadrat to the year when probability of establishment of L. dispar was greater than 99% (i.e., waiting time to establishment after first detection). To assess the effects of environmental and anthropogenic variables on each quadrat's waiting time to establishment, we performed linear mixed-effects regression models for the full transition zone and three subregions within the zone. Seasonal temperatures were found to be the primary drivers of local establishment rates. Winter temperatures had the strongest effects, especially in the northern parts of the transition zone. Furthermore, the effects of some factors on waiting times to establishment varied across subregions. Our findings contribute to identifying especially vulnerable areas to further L. dispar spread and informing region-specific criteria by invasion managers for the prioritization of areas for treatment.

The spatial synchrony of species richness and its relationship to ecosystem stability.

Synchrony is broadly important to population and community dynamics due to its ubiquity and implications for extinction dynamics, system stability, and species diversity. Investigations of synchrony in community ecology have tended to focus on covariance in the abundances of multiple species in a single location. Yet, the importance of regional environmental variation and spatial processes in community dynamics suggests that community properties, such as species richness, could fluctuate synchronously across patches in a metacommunity, in an analog of population spatial synchrony. Here, we test the prevalence of this phenomenon and the conditions under which it may occur using theoretical simulations and empirical data from 20 marine and terrestrial metacommunities. Additionally, given the importance of biodiversity for stability of ecosystem function, we posit that spatial synchrony in species richness is strongly related to stability. Our findings show that metacommunities often exhibit spatial synchrony in species richness. We also found that richness synchrony can be driven by environmental stochasticity and dispersal, two mechanisms of population spatial synchrony. Richness synchrony also depended on community structure, including species evenness and beta diversity. Strikingly, ecosystem stability was more strongly related to richness synchrony than to species richness itself, likely because richness synchrony integrates information about community processes and environmental forcing. Our study highlights a new approach for studying spatiotemporal community dynamics and emphasizes the spatial dimensions of community dynamics and stability.

Evaluating the success of treatments that slow spread of an invasive insect pest.

BACKGROUND: Treatments for the suppression and eradication of insect populations undergo substantial testing to ascertain their efficacy and safety, but the generally limited spatial and temporal scope of such studies limit knowledge of how contextual factors encountered in operational contexts shape the relative success of pest management treatments. These contextual factors potentially include ecological characteristics of the treated area, or the timing of treatments relative to pest phenology and weather events. We used an extensive database on over 1000 treatments of nascent populations of Lymantria dispar (L.) (gypsy moth) to examine how place-based and time-varying conditions shape the success of management treatments. RESULTS: We found treatment success to vary across states and years, and to be highest in small treatment blocks that are isolated from other populations. In addition, treatment success tended to be lower in treatment blocks with open forest canopies, possibly owing to challenges of effectively distributing treatments in these areas. CONCLUSIONS: Our findings emphasize the importance of monitoring for early detection of nascent gypsy moth colonies in order to successfully slow the spread of the invasion. Additionally, operations research should address best practices for effectively treating with patchy and open forest canopies. © 2021 Society of Chemical Industry.

Global hotspots of coherent marine fishery catches.

Although different fisheries can be tightly linked to each other by human and ecosystem processes, they are often managed independently. Synchronous fluctuations among fish populations or fishery catches can destabilize ecosystems and economies, respectively, but the degree of synchrony around the world remains unclear. We analyzed 1,092 marine fisheries catch time series over 60 yr to test for the presence of coherence, a form of synchrony that allows for phase-lagged relationships. We found that nearly every fishery was coherent with at least one other fishery catch time series globally and that coherence was strongest in the northeast Atlantic, western central Pacific, and eastern Indian Ocean. Analysis of fish biomass and fishing mortality time series from these hotspots revealed that coherence in biomass or fishing mortality were both possible, though biomass coherence was more common. Most of these relationships were synchronous with no time lags, and across catches in all regions, synchrony was a better predictor of regional catch portfolio effects than catch diversity. Regions with higher synchrony had lower stability in aggregate fishery catches, which can have negative consequences for food security and economic wealth.

Synchronous effects produce cycles in deer populations and deer-vehicle collisions.

Population cycles are fundamentally linked with spatial synchrony, the prevailing paradigm being that populations with cyclic dynamics are easily synchronised. That is, population cycles help give rise to spatial synchrony. Here we demonstrate this process can work in reverse, with synchrony causing population cycles. We show that timescale-specific environmental effects, by synchronising local population dynamics on certain timescales only, cause major population cycles over large areas in white-tailed deer. An important aspect of the new mechanism is specificity of synchronising effects to certain timescales, which causes local dynamics to sum across space to a substantial cycle on those timescales. We also demonstrate, to our knowledge for the first time, that synchrony can be transmitted not only from environmental drivers to populations (deer), but also from there to human systems (deer-vehicle collisions). Because synchrony of drivers may be altered by climate change, changes to population cycles may arise via our mechanism.

Self-organizing cicada choruses respond to the local sound and light environment.

Periodical cicadas exhibit an extraordinary capacity for self-organizing spatially synchronous breeding behavior. The regular emergence of periodical cicada broods across the United States is a phenomenon of longstanding public and scientific interest, as the cicadas of each brood emerge in huge numbers and briefly dominate their ecosystem. During the emergence, the 17-year periodical cicada species Magicicada cassini is found to form synchronized choruses, and we investigated their chorusing behavior from the standpoint of spatial synchrony.Cicada choruses were observed to form in trees, calling regularly every five seconds. In order to determine the limits of this self-organizing behavior, we set out to quantify the spatial synchronization between cicada call choruses in different trees, and how and why this varies in space and time.We performed 20 simultaneous recordings in Clinton State Park, Kansas, in June 2015 (Brood IV), with a team of citizen-science volunteers using consumer equipment (smartphones). We use a wavelet approach to show in detail how spatially synchronous, self-organized chorusing varies across the forest.We show how conditions that increase the strength of audio interactions between cicadas also increase the spatial synchrony of their chorusing. Higher forest canopy light levels increase cicada activity, corresponding to faster and higher-amplitude chorus cycling and to greater synchrony of cycles across space. We implemented a relaxation-oscillator-ensemble model of interacting cicadas, finding that a tendency to call more often, driven by light levels, results in all these effects.Results demonstrate how the capacity to self-organize in ecology depends sensitively on environmental conditions. Spatially correlated modulation of cycling rate by an external driver can also promote self-organization of phase synchrony.

Population spatial synchrony enhanced by periodicity and low detuning with environmental forcing.

Explaining why fluctuations in abundances of spatially disjunct populations often are correlated through time is a major goal of population ecologists. We address two hypotheses receiving little to no testing in wild populations: (i) that population cycling facilitates synchronization given weak coupling among populations, and (ii) that the ability of periodic external forces to synchronize oscillating populations is a function of the mismatch in timescales (detuning) between the force and the population. Here, we apply new analytical methods to field survey data on gypsy moth outbreaks. We report that at timescales associated with gypsy moth outbreaks, spatial synchrony increased with population periodicity via phase locking. The extent to which synchrony in temperature and precipitation influenced population synchrony was associated with the degree of mismatch in dominant timescales of oscillation. Our study provides new empirical methods and rare empirical evidence that population cycling and low detuning can promote population spatial synchrony.

Proximate determinants of Taylor's law slopes.

Taylor's law (TL), a commonly observed and applied pattern in ecology, describes variances of population densities as related to mean densities via log(variance) = log(a) + b*log(mean). Variations among datasets in the slope, b, have been associated with multiple factors of central importance in ecology, including strength of competitive interactions and demographic rates. But these associations are not transparent, and the relative importance of these and other factors for TL slope variation is poorly studied. TL is thus a ubiquitously used indicator in ecology, the understanding of which is still opaque. The goal of this study was to provide tools to help fill this gap in understanding by providing proximate determinants of TL slopes, statistical quantities that are correlated to TL slopes but are simpler than the slope itself and are more readily linked to ecological factors. Using numeric simulations and 82 multi-decadal population datasets, we here propose, test and apply two proximate statistical determinants of TL slopes which we argue can become key tools for understanding the nature and ecological causes of TL slope variation. We find that measures based on population skewness, coefficient of variation and synchrony are effective proximate determinants. We demonstrate their potential for application by using them to help explain covariation in slopes of spatial and temporal TL (two common types of TL). This study provides tools for understanding TL, and demonstrates their usefulness.

Transient synchrony among populations of five foliage-feeding Lepidoptera.

Studies of transient population dynamics have largely focused on temporal changes in dynamical behaviour, such as the transition between periods of stability and instability. This study explores a related dynamic pattern, namely transient synchrony during a 49-year period among populations of five sympatric species of forest insects that share host tree resources. The long time series allows a more comprehensive exploration of transient synchrony patterns than most previous studies. Considerable variation existed in the dynamics of individual species, ranging from periodic to aperiodic. We used time-averaged methods to investigate long-term patterns of synchrony and time-localized methods to detect transient synchrony. We investigated transient patterns of synchrony between species and related these to the species' varying density dependence structures; even species with very different density dependence exhibited at least temporary periods of synchrony. Observed periods of interspecific synchrony may arise from interactions with host trees (e.g., induced host defences), interactions with shared natural enemies or shared impacts of environmental stochasticity. The transient nature of synchrony observed here raises questions both about the identity of synchronizing mechanisms and how these mechanisms interact with the endogenous dynamics of each species. We conclude that these patterns are the result of interspecific interactions that act only temporarily to synchronize populations, after which differences in the endogenous population dynamics among the species acts to desynchronize their dynamics.