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USGS2020

Ecological prediction at macroscales using big data: Does sampling design matter?

Although ecosystems respond to global change at regional to continental scales (i.e., macroscales), model predictions of ecosystem responses often rely on data from targeted monitoring of a small proportion of sampled ecosystems within a particular geographic area. In this study, we examined how the sampling strategy used to collect data for such models influences predictive performance. We subsampled a large and spatially-extensive dataset to investigate how macroscale sampling strategy affects prediction of ecosystem characteristics in 6,784 lakes across a 1.8 million km2 area. We estimated model predictive performance for different subsets of the dataset to mimic three common sampling strategies for collecting observations of ecosystem characteristics: random sampling design, stratified random sampling design, and targeted sampling. We found that sampling strategy influenced model predictive performance such that (1) stratified random sampling designs did not improve predictive performance compared to simple random sampling designs and (2) although one of the scenarios that mimicked targeted (non-random) sampling had the poorest performing predictive models, the other targeted sampling scenarios resulted in models with similar predictive performance to that of the random sampling scenarios. Our results suggest that although potential biases in datasets from some forms of targeted sampling may limit predictive performance, compiling existing spatially-extensive datasets can result in models with good predictive performance that may inform a wide range of science questions and policy goals related to global change.

Ecological Applications↗
USGS2020

Optimal spatial prioritization of control resources for elimination of invasive species under demographic uncertainty

Populations of invasive species often spread heterogeneously across a landscape, consisting of local populations that cluster in space but are connected by dispersal. A fundamental dilemma for invasive species control is how to optimally allocate limited fiscal resources across local populations. Theoretical work based on perfect knowledge of demographic connectivity suggests that targeting local populations from which migrants originate (sources) can be optimal. However, demographic processes such as abundance and dispersal can be highly uncertain, and the relationship between local population density and damage costs (damage function) is rarely known. We used a metapopulation model to understand how budget and uncertainty in abundance, connectivity, and the damage function, together impact return on investment (ROI) for optimal control strategies. Budget, observational uncertainty, and the damage function had strong effects on the optimal resource allocation strategy. Uncertainty in dispersal probability was the least important determinant of ROI. The damage function determined which resource prioritization strategy was optimal when connectivity was symmetric but not when it was asymmetric. When connectivity was asymmetric, prioritizing source populations had a higher ROI than allocating effort equally across local populations, regardless of the damage function, but uncertainty in connectivity structure and abundance reduced ROI of the optimal prioritization strategy by 57% on average depending on the control budget. With low budgets (monthly removal rate of 6.7% of population), there was little advantage to prioritizing resources, especially when connectivity was high or symmetric, and observational uncertainty had only minor effects on ROI. Allotting funding for improved monitoring appeared to be most important when budgets were moderate (monthly removal of 13–20% of the population). Our result showed that multiple sources of observational uncertainty should be considered concurrently for optimizing ROI. Accurate estimates of connectivity direction and abundance were more important than accurate estimates of dispersal rates. Developing cost-effective surveillance methods to reduce observational uncertainties, and quantitative frameworks for determining how resources should be spatially apportioned to multiple monitoring and control activities are important and challenging future directions for optimizing ROI for invasive species control programs.

Ecological Applications↗
USGS2020

Model selection for the North American Breeding Bird Survey

The North American Breeding Bird Survey (BBS) provides data that can be used in complex, multiscale analyses of population change, while controlling for scale‐specific nuisance factors. Many alternative models can be fit to the data, but most model selection procedures are not appropriate for hierarchical models. Leave‐one‐out cross‐validation (LOOCV), in which relative model fit is assessed by omitting an observation and assessing the prediction of a model fit using the remainder of the data, provides a reasonable approach for assessing models, but is time consuming and not feasible to apply for all observations in large data sets. We report the first large‐scale formal model selection for BBS data, applying LOOCV to stratified random samples of observations from BBS data. Our results are for 548 species of North American birds, comparing the fit of four alternative models that differ in year effect structures and in descriptions of extra‐Poisson overdispersion. We use a hierarchical model among species to evaluate posterior probabilities that models are best for individual species. Models in which differences in year effects are conditionally independent (D models) were generally favored over models in which year effects are modeled by a slope parameter and a random year effect (S models), and models in which extra‐Poisson overdispersion effects are independent and t ‐distributed (H models) tended to be favored over models where overdispersion was independent and normally distributed. Our conclusions lead us to recommend a change from the conventional S model to D and H models for the vast majority of species (544/548). Comparison of estimated population trends based on the favored model relative to the S model currently used for BBS summaries indicates no consistent differences in estimated trends. Of the 18 species that showed large differences in estimated trends between models, estimated trends from the default S model were more extreme, reflecting the influence of the slope parameter in that model for species that are undergoing large population changes. WAIC, a computationally simpler alternative to LOOCV, does not appear to be a reliable alternative to LOOCV.

Ecological Applications↗
USGS2020

A novel quantitative framework for riverscape genetics

Riverscape genetics, which applies concepts in landscape genetics to riverine ecosystems, lack appropriate quantitative methods that address the spatial autocorrelation structure of linear stream networks and account for bidirectional geneflow. To address these challenges, we present a general framework for the design and analysis of riverscape genetic studies. Our framework starts with the estimation of pairwise genetic distance at sample sites and the development of a spatially structured ecological network (SSEN) on which riverscape covariates are measured. We then introduce the novel bidirectional geneflow in riverscapes (BGR) model that uses principles of isolation-by-resistance to quantify the effects of environmental covariates on genetic connectivity, with spatial covariance defined using simultaneous autoregressive models on the SSEN and the generalized Wishart distribution to model pairwise distance matrices arising through a random walk model of geneflow. We highlight the utility of this framework in an analysis of riverscape genetics for brook trout ( Salvelinus fontinalis ) in north central Pennsylvania, USA. Using the fixation index ( F ST ) as the measure of genetic distance, we estimated the effects of 12 riverscape covariates on geneflow by evaluating the relative support of eight competing BGR models. We then compared the performance of the top-ranked BGR model to results obtained from comparable analyses using multiple regression on distance matrices (MRM) and the program STRUCTURE. We found that the BGR model had more power to detect covariate effects, particularly for variables that were only partial barriers to geneflow and/or uncommon in the riverscape, making it more informative for assessing patterns of population connectivity and identifying threats to species conservation. This case study highlights the utility of our modeling framework over other quantitative methods in riverscape genetics, particularly the ability to rigorously test hypotheses about factors that influence geneflow and probabilistically estimate the effect of riverscape covariates, including stream flow direction. This framework is flexible across taxa and riverine networks, is easily executable, and provides intuitive results that can be used to investigate the likely outcomes of current and future management scenarios.

Ecological Applications↗
USGS2020

Quantifying drought’s influence on moist soil seed vegetation in California’s Central Valley through time-series remote sensing

Californias Central Valley, USA is a critical component of the Pacific Flyway despite loss of more than 90% of its wetlands. Moist soil seed (MSS) wetland plants are now produced by mimicking seasonal flooding in managed wetlands to provide an essential food resource for waterfowl. Managers need MSS plant area and productivity estimates to support waterfowl conservation, yet this remains unknown at the landscape scale. Also the effects of recent drought on MSS plants have not been quantified. We generated Landsat-derived estimates of extents and productivity (seed yield or its proxy, the green chlorophyll index) of major MSS plants including watergrass (Echinochloa crusgalli) and smartweed (Polygonum spp.) (WGSW), and swamp timothy (Crypsis schoenoides) (ST) in all Central Valley managed wetlands from 20072017. We tested the effects of water year, land ownership and region on plant area and productivity with a multifactor nested analysis of variance. For the San Joaquin Valley we explored the association between water year and water supply, and we developed metrics to support management decisions. MSS plant area maps were based on a support vector machine classification of Landsat phenology metrics (2017 map overall accuracy: 89%). ST productivity maps were created with a linear regression model of seed yield (n=68, R2 = 0.53, normalized RMSE = 10.5%). The Central Valley-wide estimated area for ST in 2017 was 32,369 ha 2,524 ha (95% C.I.), and 13,012 ha 1,384 ha for WGSW. Mean ST seed yield ranged from 577 kg/ha in the Delta Basin to 365 kg/ha in the San Joaquin Basin. WGSW area and ST seed yield decreased while ST area increased in critical drought years compared to normal water years (Scheffes test, p<0.05). Greatest ST area increases occurred in the Sacramento Valley (~75%). Voluntary water deliveries increased in normal water years, and ST seed yield increased with water supply. Z-scores of ST seed yield can be used to evaluate wetland performance and aid resource allocation decisions. Updated maps will support habitat monitoring, conservation planning and water management in future years, which are likely to face greater uncertainty in water availability with climate change.

California↗
USGS2020

Forest density intensifies the importance of snowpack to growth in water-limited pine forests

Warming climate and resulting declines in seasonal snowpack have been associated with drought stress and tree mortality in seasonally snow‐covered watersheds worldwide. Meanwhile, increasing forest density has further exacerbated drought stress due to intensified tree‐tree competition. Using a uniquely detailed dataset of population‐level forest growth (n=2495 sampled trees), we examined how inter‐annual variability in growth relates to snow volume across a range of forest densities (e.g., competitive environments) in sites spanning a broad aridity gradient across the United States. Forest growth was positively related to snowpack in water‐limited forests located at low latitude, and this relationship was intensified by forest density. However, forest growth was negatively related to snowpack in a higher latitude more energy‐limited forest, and this relationship did not interact with forest density. Future reductions in snowpack may have contrasting consequences, as growth may respond positively in energy‐limited forests and negatively in water‐limited forests; however, these declines may be mitigated by reducing stand density through forest thinning.

Ecological Applications↗
USGS2020

Tracking rates of postfire conifer regeneration vs. deciduous vegetation recovery across the western United States

Postfire shifts in vegetation composition will have broad ecological impacts. However, information characterizing postfire recovery patterns and their drivers are lacking over large spatial extents. In this analysis, we used Landsat imagery collected when snow cover (SCS) was present, in combination with growing season (GS) imagery, to distinguish evergreen vegetation from deciduous vegetation. We sought to (1) characterize patterns in the rate of postfire, dual-season Normalized Difference Vegetation Index (NDVI) across the region, (2) relate remotely sensed patterns to field-measured patterns of re-vegetation, and (3) identify seasonally specific drivers of postfire rates of NDVI recovery. Rates of postfire NDVI recovery were calculated for both the GS and SCS for more than 12,500 burned points across the western United States. Points were partitioned into faster and slower rates of NDVI recovery using thresholds derived from field plot data ( n = 230) and their associated rates of NDVI recovery. We found plots with conifer saplings had significantly higher SCS NDVI recovery rates relative to plots without conifer saplings, while plots with ≥50% grass/forbs/shrubs cover had significantly higher GS NDVI recovery rates relative to plots with <50%. GS rates of NDVI recovery were best predicted by burn severity and anomalies in postfire maximum temperature. SCS NDVI recovery rates were best explained by aridity and growing degree days. This study is the most extensive effort, to date, to track postfire forest recovery across the western United States. Isolating patterns and drivers of evergreen recovery from deciduous recovery will enable improved characterization of forest ecological condition across large spatial scales.

Ecological Applications↗
USGS2020

Landscape‐scale restoration minimizes tree growth vulnerability to 21st century drought in a dry forest

Increasing aridity is a challenge for forest managers and reducing stand density to minimize competition is a recognized strategy to mitigate drought impacts on growth. In many dry forests, the most widespread and common forest management programs currently being implemented focus on restoration of historical stand structures, primarily to minimize fire risk and enhance watershed function. The implications of these restoration projects for drought vulnerability are not well understood. Here, we examined how planned restoration treatments in the Four Forests Restoration Initiative, the largest forest restoration project in the United States, would alter landscape‐scale patterns of forest growth and drought vulnerability throughout the 21 st century. Using drought‐growth relationships developed within the landscape, we considered a suite of climate and treatment scenarios and estimated average forest growth and the proportion of years with extremely low growth as a measure of vulnerability to long‐term decline. Climatic shifts projected for this landscape include higher temperatures and shifting seasonal precipitation that promotes lower soil moisture availability in the early growing season and greater hot‐dry stress, conditions negatively associated with tree growth. However, drought severity and the magnitude of future growth declines was moderated by the thinning treatments. Compared to historical conditions, proportional growth in mid‐century declines by ~40% if thinning ceases or continues at the status quo pace. By comparison, proportional growth declines by only 20% if the Four Forest Restoration Initiative treatments are fully implemented, and < 10% if stands are thinned even more intensively than currently planned. Furthermore, restoration treatments resulted in dramatically fewer years with extremely low growth in the future, a recognized precursor to forest decline and eventual tree mortality. Benefits from density reduction for mitigating drought‐induced growth declines are more apparent in mid‐century and under RCP4.5 than under RCP8.5 at the end of the century. Future climate is inherently uncertain, and our results only reflect the climate projections from the representative suite of models examined. Nevertheless, these results indicate that forest restoration projects designed for other objectives also have substantial benefits for minimizing future drought vulnerability in dry forests and provide additional incentive to accelerate the pace of restoration.

Ecological Applications↗
USGS2021

Resistance and resilience of pelagic and littoral fishes to drought in the San Francisco Estuary

Many estuarine ecosystems and the fish communities that inhabit them have undergone substantial changes in the past several decades, largely due to multiple interacting stressors that are often of anthropogenic origin. Few are more impactful than droughts, which are predicted to increase in both frequency and severity with climate change. In this study, we examined over five decades of fish monitoring data from the San Francisco Estuary, California, USA, to evaluate the resistance and resilience of fish communities to disturbance from prolonged drought events. High resistance was defined by the lack of decline in species occurrence from a wet to a subsequent drought period, while high resilience was defined by the increase in species occurrence from a drought to a subsequent wet period. We found some unifying themes connecting the multiple drought events over the 50‐yr period. Pelagic fishes consistently declined during droughts (low resistance), but exhibit a considerable amount of resiliency and often rebound in the subsequent wet years. However, full recovery does not occur in all wet years following droughts, leading to permanently lower baseline numbers for some pelagic fishes over time. In contrast, littoral fishes seem to be more resistant to drought and may even increase in occurrence during dry years. Based on the consistent detrimental effects of drought on pelagic fishes within the San Francisco Estuary and the inability of these fish populations to recover in some years, we conclude that freshwater flow remains a crucial but not sufficient management tool for the conservation of estuarine biodiversity.

California↗
USGS2021

A Bayesian Dirichlet process community occupancy model to estimate community structure and species similarity

Community occupancy models estimate species‐specific parameters while sharing information across species by treating parameters as sampled from a common distribution. When communities consist of discrete groups, shrinkage of estimates towards the community mean can mask differences among groups. Infinite mixture models using a Dirichlet process (DP) distribution, in which the number of latent groups is estimated from the data, have been proposed as a solution. In addition to community structure, these models estimate species similarity, which allows testing hypotheses about whether traits drive species response to environmental conditions. We develop a community occupancy model (COM) using a DP distribution to model species‐level parameters. Because clustering algorithms are sensitive to dimensionality and distinctiveness of clusters, we conducted a simulation study to explore performance of the DP‐COM with different dimensions (i.e., different numbers of model parameters with species‐level DP random effects) and under varying cluster differences. Because the DP‐COM is computationally expensive, we compared its estimates to a COM with a normal random species effect. We further applied the DP‐COM model to a bird dataset from Uganda. Estimates of the number of clusters and species cluster identity improved with increasing difference among clusters and increasing dimensions of the DP; but the number of clusters was always overestimated. Estimates of number of sites occupied and species and community level covariate coefficients on occupancy probability were generally unbiased with (near‐) nominal 95% Bayesian Credible Interval coverage. Accuracy of estimates from the normal and the DP‐COM were similar. The DP‐COM clustered 166 bird species into 27 clusters regarding their affiliation with open or woodland habitat and distance to oil wells. Estimates of covariate coefficients were similar between a normal and the DP‐COM. Except sunbirds, species within a family were not more similar in their response to these covariates than the overall community. Given that estimates were consistent between the normal and the DP‐COM, and considering the computational burden for the DP models, we recommend using the DP‐COM only when the analysis focuses on community structure and species similarity, as these quantities can only be obtained under the DP‐COM.

Ecological Applications↗
USGS2021

Effects of snake fungal disease on short‐term survival, behavior, and movement in free‐ranging snakes

Pathogenic fungi are increasingly associated with epidemics in wildlife populations. Snake fungal disease (SFD, also referred to as Ophidiomycosis) is an emerging threat to snakes, taxa that are elusive and difficult to sample. Thus, assessments of the effects of SFD on populations have rarely occurred. We used a field technique to enhance detection, Passive Integrated Transponder (PIT) telemetry, and a multi‐state capture–mark–recapture model to assess SFD effects on short‐term (within‐season) survival, movement, and surface activity of two wild snake species, Regina septemvittata (Queensnake) and Nerodia sipedon (Common Watersnake). We were unable to detect an effect of disease state on short‐term survival for either species. However, we estimated Bayesian posterior probabilities of >0.99 that R. septemvittata with SFD spent more time surface‐active and were less likely to permanently emigrate from the study area. We also estimated probabilities of 0.98 and 0.87 that temporary immigration and temporary emigration rates, respectively, were lower in diseased R. septemvittata . We found evidence of elevated surface activity and lower temporary immigration rates in diseased N. sipedon , with estimated probabilities of 0.89, and found considerably less support for differences in permanent or temporary emigration rates. This study is the first to yield estimates for key demographic and behavioral parameters (survival, emigration, surface activity) of snakes in wild populations afflicted with SFD. Given the increase in surface activity of diseased snakes, future surveys of snake populations could benefit from exploring longer‐term demographic consequences of SFD and recognize that disease prevalence in surface‐active animals may exceed that of the population as a whole.

Ecological Applications↗
USGS2021

Increasing comparability among coral bleaching experiments

Coral bleaching is the single largest global threat to coral reefs worldwide. Integrating the diverse body of work on coral bleaching is critical to understanding and combating this global problem. Yet investigating the drivers, patterns, and processes of coral bleaching poses a major challenge. A recent review of published experiments revealed a wide range of experimental variables used across studies. Such a wide range of approaches enhances discovery, but without full transparency in the experimental and analytical methods used, can also make comparisons among studies challenging. To increase comparability but not stifle innovation, we propose a common framework for coral bleaching experiments that includes consideration of coral provenance, experimental conditions, and husbandry. For example, reporting the number of genets used, collection site conditions, the experimental temperature offset(s) from the maximum monthly mean (MMM) of the collection site, experimental light conditions, flow, and the feeding regime will greatly facilitate comparability across studies. Similarly, quantifying common response variables of endosymbiont (Symbiodiniaceae) and holobiont phenotypes (i.e., color, chlorophyll, endosymbiont cell density, mortality, and skeletal growth) could further facilitate cross-study comparisons. While no single bleaching experiment can provide the data necessary to determine global coral responses of all corals to current and future ocean warming, linking studies through a common framework as outlined here, would help increase comparability among experiments, facilitate synthetic insights into the causes and underlying mechanisms of coral bleaching, and reveal unique bleaching responses among genets, species, and regions. Such a collaborative framework that fosters transparency in methods used would strengthen comparisons among studies that can help inform coral reef management and facilitate conservation strategies to mitigate coral bleaching worldwide.

Ecological Applications↗
USGS2021

Evaluating natural experiments in ecology: Using synthetic controls in assessments of remotely sensed land treatments

Many important ecological phenomena occur on large spatial scales and/or are unplanned and thus do not easily fit within analytical frameworks that rely on randomization, replication, and interspersed a priori controls for statistical comparison. Analyses of such large‐scale, natural experiments are common in the health and econometrics literature, where techniques have been developed to derive insight from large, noisy observational data sets. Here, we apply a technique from this literature, synthetic control, to assess landscape change with remote sensing data. The basic data requirements for synthetic control include (1) a discrete set of treated and untreated units, (2) a known date of treatment intervention, and (3) time series response data that include both pre‐ and post‐treatment outcomes for all units. Synthetic control generates a response metric for treated units relative to a no‐action alternative based on prior relationships between treated and unexposed groups. Using simulations and a case study involving a large‐scale brush‐clearing management event, we show how synthetic control can intuitively infer treatment effect sizes from satellite data, even in the presence of confounding noise from climate anomalies, long‐term vegetation dynamics, or sensor errors. We find that accuracy depends on the number and quality of potential control units, highlighting the importance of selecting appropriate control populations. Although we consider the synthetic control approach in the context of natural experiments with remote sensing data, we expect the methodology to have wider utility in ecology, particularly for systems with large, complex, and poorly replicated experimental units.

Ecological Applications↗
USGS2021

Time-to-detection occupancy methods: Performance and utility for improving efficiency of surveys

Occupancy methods propelled the quantitative study of species distributions forward by separating the observation process, or the imperfect detectability of species, from the ecological processes of interest governing species distributions. Occupancy studies come at a cost, however: the collection of additional data to account for nondetections at sites where the species is present. The most common occupancy designs (repeated measures designs) require repeat visits to sites or the use of multiple observers or detection methods. Time‐to‐detection methods have been identified as a potentially efficient alternative, requiring only one visit to each site by a single observer. A comparison of time‐to‐detection methods to repeated measures designs for visual encounter surveys would allow researchers to evaluate whether time‐to‐detection methods might be appropriate for their study system and can inform optimal survey design. We collected time‐to‐detection data during two different repeated measures design occupancy surveys for four amphibians and compared the performance of time‐to‐detection methods to the other designs using the location (potential bias) and precision of posterior distributions for occurrence parameters. We further used results of time‐to‐detection surveys to optimize survey design. Time‐to‐detection methods performed best for species that are widespread and have high detection probabilities and rates, but performed less well for cryptic species with lower probability of occurrence or whose detection was strongly affected by survey conditions. In all cases single surveys were most efficient in terms of person‐hours expended, but under some conditions the survey duration required to achieve high detection probabilities would be prohibitively long for a single survey. Regardless of occupancy survey design, time‐to‐detection methods provide important information that can be used to optimize surveys, allowing researchers and resource managers to efficiently achieve monitoring and conservation goals. Collecting time‐to‐detection data while conducting repeated measures occupancy surveys requires only small modifications to field methods but could have large benefits in terms of time spent surveying in the long‐term.

California↗
USGS2021

Water storage decisions will determine the distribution and persistence of imperiled river fishes

Managing the world’s freshwater supply to meet societal and environmental needs in a changing climate is one of the biggest challenges for the 21 st century. Dams provide water security, however, the allocation of dwindling water supply among reservoirs could exacerbate or ameliorate the effects of climate change on aquatic communities. Here, we show that the relative sensitivity of river thermal regimes to direct impacts of climate change and societal decisions concerning water storage vary substantially throughout a river basin. In the absence of interspecific interactions, future Colorado River temperatures would appear to benefit both endemic and nonnative fish species. However, endemic species are already declining or extirpated in locations where their ranges overlap with warmwater nonnatives and changes in water storage may lead to warming in some of the coolest portions of the river basin, facilitating further nonnative expansion. Integrating environmental considerations into ongoing water storage negotiations may lead to better resource outcomes than mitigating nonnative species impacts after the fact.

Ecological Applications↗
USGS2021

Effects of postfire climate and seed availability on postfire conifer regeneration

Large, severe fires are becoming more frequent in many forest types across the western United States and have resulted in tree mortality across tens of thousands of hectares. Conifer regeneration in these areas is limited because seeds must travel long distances to reach the interior of large burned patches and establishment is jeopardized by increasingly hot and dry conditions. To better inform postfire management in low elevation forests of California, USA, we collected 5‐year postfire recovery data from 1,234 study plots in 19 wildfires that burned from 2004–2012 and 18 years of seed production data from 216 seed fall traps (1999–2017). We used this data in conjunction with spatially extensive estimates of climate, topography, forest composition, and burn severity to construct taxon‐specific, spatially explicit models of conifer regeneration that incorporate estimated climate conditions and seed availability during postfire recovery windows. We found that after accounting for other predictors both postfire and historical precipitation were strong predictors of regeneration, suggesting that both direct effects of postfire moisture conditions and biological inertia from historical climate may play a role in regeneration. Alternatively, postfire regeneration may simply be driven by postfire climate and apparent relationships with historical climate could be spurious. The estimated sensitivity of regeneration to postfire seed availability was strongest in firs and all conifers combined and weaker in pines. Seed production exhibited high temporal variability with seed production varying by over two orders of magnitude among years. Our models indicate that during droughts postfire conifer regeneration declines most substantially in low‐to‐moderate elevation forests. These findings enhance our mechanistic understanding of forecasted and historically documented shifts in the distribution of trees.

California↗
USGS2021

Life-history theory provides a framework for detecting resource limitation: A test of the Nutritional Buffer Hypothesis

For ungulates and other long-lived species, life-history theory predicts that nutritional reserves are allocated to reproduction in a state-dependent manner because survival is highly conserved. Further, as per capita food abundance and nutritional reserves decline (i.e., density dependence intensifies), reproduction and recruitment become increasingly sensitive to weather. Thus, the degree to which weather influences vital rates should be associated with proximity to nutritional carrying capacity—a notion that we refer to as the Nutritional Buffer Hypothesis. We tested the Nutritional Buffer Hypothesis using six moose ( Alces alces ) populations that varied in calf recruitment (33–69 calves/100 cows). We predicted that populations with high calf recruitment were nutritionally buffered against the effects of unfavorable weather, and thus were below nutritional carrying capacity. We applied a suite of tools to quantify habitat and nutritional condition of each population and found that increased browse condition, forage quality, and body fat were associated with increased pregnancy and calf recruitment, thereby providing multiple lines of evidence that declines in calf recruitment were underpinned by resource limitation. From 2001 to 2015, recruitment was more sensitive to interannual variation in weather (e.g., winter severity, drought) and plant phenology (e.g., duration of spring) for populations with reduced browse condition, forage quality, and body fat, suggesting these populations lacked the nutritional reserves necessary to buffer demographic performance against the effects of unfavorable weather. Further, average within-population calf recruitment was determined by regional climatic variation, suggesting that the pattern of reduced recruitment near the southern range boundary of moose stems from an interaction between climate and resource limitation. When coupled with information on habitat, nutrition, weather, and climate, life-history theory provides a framework to estimate nutritional limitation, proximity to nutritional carrying capacity, and impacts of climate change for ungulates.

Colorado, Utah, Wyoming↗