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Moving forward in circles: Challenges and opportunities in modeling population cycles

Population cycling is a widespread phenomenon, observed across a multitude of taxa in both laboratory and natural conditions. Historically, the theory associated with population cycles was tightly linked to pairwise consumer–resource interactions and studied via deterministic models, but current empirical and theoretical research reveals a much richer basis for ecological cycles. Stochasticity and seasonality can modulate or create cyclic behaviour in non-intuitive ways, the high-dimensionality in ecological systems can profoundly influence cycling, and so can demographic structure and eco-evolutionary dynamics. An inclusive theory for population cycles, ranging from ecosystem-level to demographic modelling, grounded in observational or experimental data, is therefore necessary to better understand observed cyclical patterns. In turn, by gaining better insight into the drivers of population cycles, we can begin to understand the causes of cycle gain and loss, how biodiversity interacts with population cycling, and how to effectively manage wildly fluctuating populations, all of which are growing domains of ecological research.

Ecology Letters

Carrying capacity in a heterogeneous environment with habitat connectivity

A large body of theory predicts that populations diffusing in heterogeneous environments reach higher total size than if non-diffusing, and, paradoxically, higher size than in a corresponding homogeneous environment. However, this theory and its assumptions have not been rigorously tested. Here, we extended previous theory to include exploitable resources, proving qualitatively novel results, which we tested experimentally using spatially diffusing laboratory populations of yeast. Consistent with previous theory, we predicted and experimentally observed that spatial diffusion increased total equilibrium population abundance in heterogeneous environments, with the effect size depending on the relationship between r and K . Refuting previous theory, however, we discovered that homogeneously distributed resources support higher total carrying capacity than heterogeneously distributed resources, even with species diffusion. Our results provide rigorous experimental tests of new and old theory, demonstrating how the traditional notion of carrying capacity is ambiguous for populations diffusing in spatially heterogeneous environments.

Ecology Letters

Making ecological models adequate

Critical evaluation of the adequacy of ecological models is urgently needed to enhance their utility in developing theory and enabling environmental managers and policymakers to make informed decisions. Poorly supported management can have detrimental, costly or irreversible impacts on the environment and society. Here, we examine common issues in ecological modelling and suggest criteria for improving modelling frameworks. An appropriate level of process description is crucial to constructing the best possible model, given the available data and understanding of ecological structures. Model details unsupported by data typically lead to over parameterisation and poor model performance. Conversely, a lack of mechanistic details may limit a model's ability to predict ecological systems’ responses to management. Ecological studies that employ models should follow a set of model adequacy assessment protocols that include: asking a series of critical questions regarding state and control variable selection, the determinacy of data, and the sensitivity and validity of analyses. We also need to improve model elaboration, refinement and coarse graining procedures to better understand the relevancy and adequacy of our models and the role they play in advancing theory, improving hind and forecasting, and enabling problem solving and management.

Ecology Letters

Artificial light at night confounds broad-scale habitat use by migrating birds

With many of the world's migratory bird populations in alarming decline, broad-scale assessments of responses to migratory hazards may prove crucial to successful conservation efforts. Most birds migrate at night through increasingly light-polluted skies. Bright light sources can attract airborne migrants and lead to collisions with structures, but might also influence selection of migratory stopover habitat and thereby acquisition of food resources. We demonstrate, using multi-year weather radar measurements of nocturnal migrants across the northeastern U.S., that autumnal migrant stopover density increased at regional scales with proximity to the brightest areas, but decreased within a few kilometers of brightly-lit sources. This finding implies broad-scale attraction to artificial light while airborne, impeding selection for extensive forest habitat. Given that high-quality stopover habitat is critical to successful migration, and hindrances during migration can decrease fitness, artificial lights present a potentially heightened conservation concern for migratory bird populations.

Connecticut, Delaware, Maine, Maryland, Massachuse

Ecological genomics predicts climate vulnerability in an endangered southwestern songbird

Few regions have been more severely impacted by climate change in the USA than the Desert Southwest. Here, we use ecological genomics to assess the potential for adaptation to rising global temperatures in a widespread songbird, the willow flycatcher ( Empidonax traillii ), and find the endangered desert southwestern subspecies ( E. t. extimus ) most vulnerable to future climate change. Highly significant correlations between present abundance and estimates of genomic vulnerability – the mismatch between current and predicted future genotype–environment relationships – indicate small, fragmented populations of the southwestern willow flycatcher will have to adapt most to keep pace with climate change. Links between climate‐associated genotypes and genes important to thermal tolerance in birds provide a potential mechanism for adaptation to temperature extremes. Our results demonstrate that the incorporation of genotype–environment relationships into landscape‐scale models of climate vulnerability can facilitate more precise predictions of climate impacts and help guide conservation in threatened and endangered groups.

Ecology Letters

Running on empty: Recharge dynamics from animal movement data

Vital rates such as survival and recruitment have always been important in the study of population and community ecology. At the individual level, physiological processes such as energetics are critical in understanding biomechanics and movement ecology and also scale up to influence food webs and trophic cascades. Although vital rates and population-level characteristics are tied with individual-level animal movement, most statistical models for telemetry data are not equipped to provide inference about these relationships because they lack the explicit, mechanistic connection to physiological dynamics. We present a framework for modelling telemetry data that explicitly includes an aggregated physiological process associated with decision making and movement in heterogeneous environments. Our framework accommodates a wide range of movement and physiological process specifications. We illustrate a specific model formulation in continuous-time to provide direct inference about gains and losses associated with physiological processes based on movement. Our approach can also be extended to accommodate auxiliary data when available. We demonstrate our model to infer mountain lion ( Puma concolor ; in Colorado, USA) and African buffalo ( Syncerus caffer ; in Kruger National Park, South Africa) recharge dynamics.

Ecology Letters

Non-native plants have greater impacts because of differing per-capita effects and non-linear abundance-impact curves

Invasive, non-native species can have tremendous impacts on biotic communities, where they reduce the abundance and diversity of local species. However, it remains unclear whether impacts of non-native species arise from their high abundance or whether each non-native individual has a disproportionate impact – i.e., a higher per-capita effect – on co-occurring species compared to impacts by native species. Using a long-term study of wetlands, we asked how temporal variation in dominant native and non-native plants impacted the abundance and richness of other plants in the recipient community. Non-native plants reached higher abundances than natives and had greater per-capita effects. The abundance-impact relationship between plant abundance and richness was nonlinear. Compared with increasing native abundance, increasing non-native abundance was associated with steeper declines in richness because of greater per-capita effects and nonlinearities in the abundance-impact relationship. Our study supports eco-evolutionary novelty of non-natives as a driver of their outsized impacts on communities.

Illinois

Transient population dynamics impede restoration and may promote ecosystem transformation after disturbance

The apparent failure of ecosystems to recover from increasingly widespread disturbance is a global concern. Despite growing focus on factors inhibiting resilience and restoration, we still know very little about how demographic and population processes influence recovery. Using inverse and forward demographic modelling of 531 post‐fire sagebrush populations across the western US, we show that demographic processes during recovery from seeds do not initially lead to population growth but rather to years of population decline, low density, and risk of extirpation after disturbance and restoration, even at sites with potential to support long‐term, stable populations. Changes in population structure, and resulting transient population dynamics, lead to a > 50% decline in population growth rate after disturbance and significant reductions in population density. Our results indicate that demographic processes influence the recovery of ecosystems from disturbance and that demographic analyses can be used by resource managers to anticipate ecological transformation risk.

Ecology Letters

Land-use change and the ecological consequences of personality in small mammals

Many plants rely on animals for seed dispersal, but are all individuals equally effective at dispersing seeds? If not, then the loss of certain individual dispersers from populations could have cascade effects on ecosystems. Despite the importance of seed dispersal for forest ecosystems, variation among individual dispersers and whether land‐use change interferes with this process remains untested. Through a large‐scale field experiment conducted on small mammal seed dispersers, we show that an individual's personality affects its choice of seeds, as well as how distant and where seeds are cached. We also show that anthropogenic habitat modifications shift the distribution of personalities within a population, by increasing the proportion of bold, active, and anxious individuals and in‐turn affecting the potential survival and dispersal of seeds. We demonstrate that preserving diverse personality types within a population is critical for maintaining the key ecosystem function of seed dispersal.

Ecology Letters

Spatial and temporal variation of ecosystem properties at macroscales

Although spatial and temporal variation in ecological properties has been well-studied, crucial knowledge gaps remain for studies conducted at macroscales and for ecosystem properties related to material and energy. We test four propositions of spatial and temporal variation in ecosystem properties within a macroscale (1000 km's) extent. We fit Bayesian hierarchical models to thousands of observations from over two decades to quantify four components of variation – spatial (local and regional) and temporal (local and coherent); and to model their drivers. We found strong support for three propositions: (1) spatial variation at local and regional scales are large and roughly equal, (2) annual temporal variation is mostly local rather than coherent, and, (3) spatial variation exceeds temporal variation. Our findings imply that predicting ecosystem responses to environmental changes at macroscales requires consideration of the dominant spatial signals at both local and regional scales that may overwhelm temporal signals.

Ecology Letters

Spatial memory shapes migration and its benefits: Evidence from a large herbivore

From fine-scale foraging to broad-scale migration, animal movement is shaped by the distribution of resources. There is mounting evidence, however, that learning and memory also guide movement. Although migratory mammals commonly track resource waves, how resource tracking and memory guide long-distance migration has not been reconciled. We examined these hypotheses using movement data from four populations of migratory mule deer ( n = 91). Spatial memory had an extraordinary influence on migration, affecting movement 2–28 times more strongly than tracking spring green-up or autumn snow depth. Importantly, with only an ability to track resources, simulated deer were unable to recreate empirical migratory routes. In contrast, simulated deer with memory of empirical routes used those routes and obtained higher foraging benefits. For migratory terrestrial mammals, spatial memory provides knowledge of where seasonal ranges and migratory routes exist, whereas resource tracking determines when to beneficially move within those areas.

Colorado, Wyoming

Migratory divides coincide with reproductive barriers across replicated avian hybrid zones above the Tibetan Plateau

Migratory divides are proposed to be catalysts for speciation across a diversity of taxa. However, it is difficult to test the relative contributions of migratory behaviour vs. other divergent traits to reproductive isolation. Comparing hybrid zones with and without migratory divides offers a rare opportunity to directly examine the contribution of divergent migratory behaviour to reproductive barriers. We show that across replicate sampling transects of two pairs of barn swallow ( Hirundo rustica ) subspecies, strong reproductive isolation coincided with a migratory divide spanning 20 degrees of latitude. A third subspecies pair exhibited no evidence for a migratory divide and hybridised extensively. Within migratory divides, overwintering habitats were associated with assortative mating, implicating a central contribution of divergent migratory behaviour to reproductive barriers. The remarkable geographic coincidence between migratory divides and genetic breaks supports a long-standing hypothesis that the Tibetan Plateau is a substantial barrier contributing to the diversity of Siberian avifauna.

Tibetan Plateau

From theory to experiments for testing the proximate mechanisms of mast seeding: An agenda for an experimental ecology

Highly variable and synchronised production of seeds by plant populations is called masting and is implicated in many important ecological processes, but how it arises remains poorly understood. The lack of experimental studies prevents underlying mechanisms from being explicitly tested, and thereby precludes meaningful predictions on the consequences of changing environments for plant reproductive patterns and global vegetation dynamics. Here we review the most relevant hypothetical drivers of masting and outline a research agenda that takes the biology of masting from a largely observational field of ecology to one rooted in mechanistic understanding. We divide the experimental framework into three main processes: resource dynamics, pollen limitation, and genetic and hormonal regulation, and illustrate how specific predictions about proximate mechanisms can be tested, highlighting the few successful experiments as examples. We envision that the experiments we outline will deliver new insights into how and why masting patterns might respond to a changing environment.

Ecology Letters

Metabolic rate is negatively linked to adult survival but does not explain latitudinal differences in songbirds

Survival rates vary dramatically among species and predictably across latitudes, but causes of this variation are unclear. The rate of living hypothesis posits that physiological damage from metabolism causes species with faster metabolic rates to exhibit lower survival rates. However, whether increased survival commonly observed in tropical and south temperate latitudes is associated with slower metabolic rate remains unclear. We compared metabolic rates and annual survival rates that we measured across 46 species, and from literature data across 147 species in northern, southern, and tropical latitudes. High metabolic rates were associated with lower survival but survival varied substantially among latitudinal regions independent of metabolism. The inability of metabolic rate to explain latitudinal variation in survival suggests 1) species may evolve physiological mechanisms that mitigate physiological damage from cellular metabolism, and 2) extrinsic rather than intrinsic sources of mortality are the primary causes of latitudinal differences in survival.

Ecology Letters

Generalizing indirect defense and resistance of plants

Indirect defence, the adaptive top‐down control of herbivores by plant traits that enhance predation, is a central component of plant–herbivore interactions. However, the scope of interactions that comprise indirect defence and associated ecological and evolutionary processes has not been clearly defined. We argue that the range of plant traits that mediate indirect defence is much greater than previously thought, and we further organise major concepts surrounding their ecological functioning. Despite the wide range of plant traits and interacting organisms involved, indirect defences show commonalities when grouped. These categories are based on whether indirect defences boost natural enemy abundance via food or shelter resources, or, alternatively, increase natural enemy foraging efficiency via information or alteration of habitat complexity. The benefits of indirect defences to natural enemies should be further explored to establish the conditions in which indirect defence generates a plant–natural enemy mutualism. By considering the broader scope of plant–herbivore–natural enemy interactions that comprise indirect defence, we can better understand plant‐based food webs, as well as the evolutionary processes that have shaped them.

Ecology Letters

Adult survival probability and body size affect parental risk-taking across latitudes

Parents faced with a predator must choose between their own safety versus taking care of their offspring. Each choice can have fitness costs. Life-history theory predicts that longer-lived species should be less willing than shorter-lived species to return to care for their offspring after a predator disturbance because they have more opportunities to reproduce in the future. We increased adult predation risk during incubation for 40 bird species in north temperate, tropical, and south temperate latitudes. We found that species with higher adult survival probabilities were more cautious, waiting longer before returning to the nest to provide care. Contrary to other studies, we also found that parents were more risk averse and waited longer to return in smaller than larger species, likely reflecting greater vulnerability of smaller species. Ultimately, the relative risk a predator poses to a species and the probability of future reproduction predict parental risk taking across the world.

Arizona, Sabah

Forest responses to last-millennium hydroclimate variability are governed by spatial variations in ecosystem sensitivity

Forecasts of future forest change are governed by ecosystem sensitivity to climate change, but ecosystem model projections are under-constrained by data at multidecadal and longer timescales. Here, we quantify ecosystem sensitivity to centennial-scale hydroclimate variability, by comparing dendroclimatic and pollen-inferred reconstructions of drought, forest composition and biomass for the last millennium with five ecosystem model simulations. In both observations and models, spatial patterns in ecosystem responses to hydroclimate variability are strongly governed by ecosystem sensitivity rather than climate exposure. Ecosystem sensitivity was higher in models than observations and highest in simpler models. Model-data comparisons suggest that interactions among biodiversity, demography and ecophysiology processes dampen the sensitivity of forest composition and biomass to climate variability and change. Integrating ecosystem models with observations from timescales extending beyond the instrumental record can better understand and forecast the mechanisms regulating forest sensitivity to climate variability in a complex and changing world.

Ecology Letters

Why disease ecology needs life-history theory: A host perspective

When facing an emerging infectious disease of conservation concern, we often have little information on the nature of the host‐parasite interaction to inform management decisions. However, it is becoming increasingly clear that the life‐history strategies of host species can be predictive of individual‐ and population‐level responses to infectious disease, even without detailed knowledge on the specifics of the host‐parasite interaction. Here, we argue that a deeper integration of life‐history theory into disease ecology is timely and necessary to improve our capacity to understand, predict and mitigate the impact of endemic and emerging infectious diseases in wild populations. Using wild vertebrates as an example, we show that host life‐history characteristics influence host responses to parasitism at different levels of organisation, from individuals to communities. We also highlight knowledge gaps and future directions for the study of life‐history and host responses to parasitism. We conclude by illustrating how this theoretical insight can inform the monitoring and control of infectious diseases in wildlife.

Ecology Letters