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Detecting resource limitation in a large herbivore population is enhanced with measures of nutritional condition

Resource limitation at the population level is a function of forage quality and its abundance relative to its per capita availability, which in turn, determines nutritional condition of individuals. Effects of resource limitation on population dynamics in ungulates often occur through predictable and sequential changes in vital rates, which can enable assessments of how resource limitation influences population growth. We tested theoretical predictions of bottom-up (i.e., resource limitation) forcing on moose ( Alces alces ) through the lens of vital rates by quantifying the relative influence of intrinsic measures of nutritional condition and extrinsic measures of remotely sensed environmental data on demographic rates. We measured rates of pregnancy, parturition, juvenile, and adult survival for 82 adult females in a population where predators largely were absent. Life stage simulation analyses (LSAs) indicated that interannual fluctuations in adult survival contributed to most of the variability in λ. We then extended the LSA to estimate vital rates as a function of bottom-up covariates to evaluate their influence on λ. We detected weak signatures of effects from environmental covariates that were remotely sensed and spatially explicit to each seasonal range. Instead, nutritional condition strongly influenced rates of pregnancy, parturition, and overwinter survival of adults, clearly implicating resource limitation on λ. Our findings depart from the classic life-history paradigm of population dynamics in ungulates in that adult survival was highly variable and generated most of the variability in population growth rates. At the surface, lack of variation explained by environmental covariates may suggest weak evidence of resource limitation in the population, when nutritional condition actually underpinned most demographics. We suggest that variability in vital rates and effects of resource limitation may depend on context more than previously appreciated, and density dependence can obfuscate the relationships between remotely sensed data and demographic rates.

Wyoming

Climate change Is likely to alter future wolf - moose - forest interactions at Isle Royale National Park, United States

We evaluated how climate change and variable rates of moose browsing intensity, as they relate to wolf predation, might affect the forests of Isle Royale National Park, Michigan, United States by conducting a modeling experiment. The experiment consisted of contrasting three different scenarios of wolf management and with a static (current conditions) and changing climate (high emissions). Our results indicate that the interactive effects of wolf predation and climate change are likely to be temporally variable and dependent on biogeographic and forest successional processes. During the first 50 years of 120-year simulations, when the effects of climate change were less impactful, higher simulated rates of predation by wolves reduced moose population densities, resulting in greater forest biomass and higher carrying capacities for moose. However, over the longer term, early successional and highly palatable aspen and birch forests transitioned to late successional spruce and fir forests, regardless of climate or predation intensity. After 50 years, the effects of climate change and predation were driven by effects on balsam fir, a late successional conifer species that is fed on by moose. High-intensity predation of moose allowed balsam fir to persist over the long term but only under the static climate scenario. The climate change scenario caused a reduction in balsam fir and the other boreal species that moose currently feed on, and the few temperate species found on this isolated island were unable to compensate for such reductions, causing strong declines in total forest biomass. The direct effects of moose population management via reintroduction of wolves may become increasingly ineffective as the climate continues to warm because the productivity of boreal plant species may not be sufficient to support a moose population, and the isolation of the island from mainland temperate tree species may reduce the likelihood of compensatory species migrations.

Michigan

Microbiomes from biorepositories? 16S rRNA bacterial amplicon sequencing of archived and contemporary intestinal samples of wild mammals (Eulipotyphla: Soricidae)

Interest in gut microbial community composition has exploded recently as a result of the increasing ability to characterize these organisms and a growing understanding of their role in host fitness. New technologies, such as next generation amplicon (16S rRNA) sequencing, have enabled identification of bacterial communities from samples of diverse origin (e.g., fecal, skin, genital, environmental, etc.). Relatively little work, however, has explored the feasibility of utilizing historical samples (e.g., museum archived samples) of varying age, quality, and preservation type. Because natural history collections span multiple decades, these biorepositories have the potential to provide fundamental historical baselines to measure and better understand biodiversity on a changing planet. Utilizing even a small proportion of museum specimens could provide a means of sampling past microbial communities, allowing for direct comparison to contemporary communities and more complete understanding of dynamic shifts through time. We examined the feasibility of obtaining 16S rRNA amplicon microbiome data from whole gastrointestinal tracts (GIs) of shrews of varying age and preservation method, including 5 freshly collected shrew GIs immediately fixed in liquid nitrogen (LN2), 10 ten-year old shrew GIs frozen at −20°C (whole animal), and 10 shrews of varying ages (4 from 1968, 1 from 1980, 1 from 2001, 1 from 2004, 1 from 2007, 1 from 2011 and 2 from 2013) fixed and stored whole in 70% ethanol. Not surprisingly, results of 16S rDNA amplicon sequencing reveal significantly different bacterial communities between different preservation techniques and age of samples. Ten-year old frozen samples had bacterial communities most similar to freshly collected (LN2) samples, while the bacterial communities of both were significantly different from the 70% ethanol preserved samples of various ages. Amongst those preserved in 70% ethanol, age of samples also influenced bacterial community composition. Additionally, we compare results of OTU based and ASV based analyses. Looking ahead, field collectors and museums should develop and adopt best practices related to frozen preservation to ensure adequate material for future microbiome investigations.

Frontiers in Ecology and Evolution

Shorebird reproductive response to exceptionally early and late springs varies across sites in Arctic Alaska

While increases in overall temperatures are widely reported in the Arctic, large inter-annual variation in spring weather, with extreme early and late conditions, is also occurring. Using data collected from three sites in Arctic Alaska, we explored how shorebird breeding density, nest initiation, nest synchrony, nest survival, and phenological mismatch varied between two exceptionally early (2015 and 2016) and late (2017 and 2018) springs. We assessed these differences in the context of long-term data from each site and whether species exhibited conservative or opportunistic reproductive strategies. Conservative shorebirds typically display nest-site fidelity and territoriality, consistent population densities, relatively even individual spacing, and monogamous mating systems with bi-parental incubation. In contrast, opportunistic shorebirds display the opposite traits, and a polygamous mating system with uniparental incubation. In this study, we evaluated 2,239 nests from 13 shorebird species, 2015–2018, and found that shorebirds of both strategies bred earlier and in higher numbers in early, warm springs relative to historic levels (based on 3,789 nests, 2005–2014); opposite trends were observed in late springs. In early springs, nests were initiated less synchronously than in late springs. Nest survival was unrelated to spring type, but was greater in earlier laid nests overall. Invertebrate food resources emerged earlier in early springs, resulting in a greater temporal asynchrony between invertebrate emergence and chick hatching in early than late springs. However, invertebrate abundance was quite variable among sites and years regardless of spring type. Overall, our results were generally consistent with predicted relationships between spring conditions and reproductive parameters. However, we detected differences among sites that could not be explained by other ecological factors (e.g., predators or alternative prey). Differences in shorebird community composition and other subtler methodological/ecological differences among sites highlight the difficulty of understanding the complex nature of these ecological systems and the importance of evaluating questions at multiple sites across multiple years. Our study demonstrates that shorebirds exhibit a high degree of behavioral flexibility in response to variable Arctic conditions, but whether this flexibility is enough to allow them to optimally track changing environmental conditions or if evolutionary adjustments will be necessary is unknown.

Alaska

Post-fire vegetation response in a repeatedly burned low-elevation sagebrush steppe protected area provides insights about resilience and invasion resistance

Sagebrush steppe ecosystems are threatened by human land-use legacies, biological invasions, and altered fire and climate dynamics. Steppe protected areas are therefore of heightened conservation importance but are few and vulnerable to the same impacts broadly affecting sagebrush steppe. To address this problem, sagebrush steppe conservation science is increasingly emphasizing a focus on resilience to fire and resistance to non-native annual grass invasion as a decision framework. It is well-established that the positive feedback loop between fire and annual grass invasion is the driving process of most contemporary steppe degradation. We use a newly developed ordinal zero-augmented beta regression model fit to large-sample vegetation monitoring data from John Day Fossil Beds National Monument, USA, spanning 7 years to evaluate fire responses of two native perennial foundation bunchgrasses and two non-native invasive annual grasses in a repeatedly burned, historically grazed, and inherently low-resilient protected area. We structured our model hierarchically to support inferences about variation among ecological site types and over time after also accounting for growing-season water deficit, fine-scale topographic variation, and burn severity. We use a state-and-transition conceptual diagram and abundances of plants listed in ecological site reference conditions to formalize our hypothesis of fire-accelerated transition to ecologically novel annual grassland. Notably, big sagebrush ( Artemisia tridentata ) and other woody species were entirely removed by fire. The two perennial grasses, bluebunch wheatgrass ( Pseudoroegneria spicata ) and Thurber's needlegrass ( Achnatherum thurberianum ) exhibited fire resiliency, with no apparent trend after fire. The two annual grasses, cheatgrass ( Bromus tectorum ) and medusahead ( Taeniatherum caput-medusae ), increased in response to burn severity, most notably medusahead. Surprisingly, we found no variation in grass cover among ecological sites, suggesting fire-driven homogenization as shrubs were removed and annual grasses became dominant. We found contrasting responses among all four grass species along gradients of topography and water deficit, informative to protected-area conservation strategies. The fine-grained influence of topography was particularly important to variation in cover among species and provides a foothold for conservation in low-resilient, aridic steppe. Broadly, our study demonstrates how to operationalize resilience and resistance concepts for protected areas by integrating empirical data with conceptual and statistical models.

Oregon

Contrasting geographic patterns of ignition probability and burn severity in the Mojave Desert

The extent and frequency of fire has increased in many arid systems over the last century, with a large proportion of area in some regions undergoing transitions to novel conditions. Portions of the Mojave Desert in southwestern North America have undergone such transitions, most often from woody to herbaceous-dominated systems. These transitions have often been attributed to the proliferation of invasive annual grasses that promote more frequent fire, but recent evidence indicates that transitions can also occur independent of fire frequency if burn severity is high. In addition, high probability of ignition (i.e., potentially high fire frequency) and high burn severity may not always be geographically related. Therefore, our goals were to: (1) map potential burn severity, fire frequency, and probability of ignition across the Mojave; and, (2) evaluate spatial association among predicted burn severity, fire frequency and probability of ignition. We first mapped perimeters of 250 wildfires > 405 ha that occurred from 1972 to 2010, then extracted data on fire frequency (number of times burned from 1972 to 2010), burn severity (the difference Normalized Burn Ratio), and 15 predictor variables representing physiography, climate, ignition, and vegetation. Maximum entropy was used to predict probability of ignition and Random Forest models were used to predict dNBR and fire frequency. Areas with high burn severity and high ignition probability had opposite spatial trends; areas with high burn severity were predicted to predominantly be in the northwest part of the region whereas areas with high ignition probability were predicted to be in the northeast. The models indicate the existence of a number of spatially structured but temporally dynamic fire regimes throughout the Mojave Desert. Two prevalent and ecologically significant regimes include one with frequent fires of low to moderate severity and another with infrequent fire of high severity. Areas with high fire frequency are currently limited in extent (<1% total area). However, cover of invasive grasses can remain high decades after a burn of high or moderate severity, so grass-fire cycles could develop in areas where there may be expectations of infrequent fire as well as those with relatively high fire frequency.

Arizona, California, Nevada, Utah

Integrating environmental DNA results with diverse data sets to improve biosurveillance of river health

Autonomous, robotic environmental (e)DNA samplers now make it possible for biological observations to match the scale and quality of abiotic measurements collected by automated sensor networks. Merging these automated data streams may allow for improved insight into biotic responses to environmental change and stressors. Here, we merged eDNA data collected by robotic samplers installed at three U.S. Geological Survey (USGS) streamgages with gridded daily weather data, and daily water quality and quantity data into a cloud-hosted database. The eDNA targets were a rare fish parasite and a more common salmonid fish. We then used computationally expedient Bayesian hierarchical occupancy models to evaluate associations between abiotic conditions and eDNA detections and to simulate how uncertainty in result interpretation changes with the frequency of autonomous robotic eDNA sample collection. We developed scripts to automate data merging, cleaning and analysis steps into a chained-step, workflow. We found that inclusion of abiotic covariates only provided improved insight for the more common salmonid fish since its DNA was more frequently detected. Rare fish parasite DNA was infrequently detected, which caused occupancy parameter estimates and covariate associations to have high uncertainty. Our simulations found that collecting samples at least once per day resulted in more detections and less parameter uncertainty than less frequent sampling. Our occupancy and simulation results together demonstrate the advantages of robotic eDNA samplers and how these samples can be combined with easy to acquire, publicly available data to foster real-time biosurveillance and forecasting.

Idaho, Wyoming, Montana

Moose habitat selection and fitness consequences during two critical winter tick life stages in Vermont, United States

The moose ( Alces alces ) is a charismatic species in decline across much of their southern distribution in North America. In the northeastern United States, much of the reduction has been attributed to winter tick ( Dermacentor albipictus ) infestations. Winter ticks are fairly immobile throughout all life stages, and therefore their distribution patterns at any given time are shaped largely by the occurrence of moose across the landscape during the peak of two critical time periods: fall questing (when ticks latch onto moose) and spring drop-off (when engorged female ticks detach from moose). We used recent land cover and lidar data within a dynamic occupancy modeling framework to estimate first-order habitat selection (use vs. non-use) of female moose ( n = 74) during the tick questing and drop-off periods. Patch extinction and colonization rates between the fall questing and spring drop-off periods were strongly influenced by habitat and elevation, but these effects were diminished during the fall questing period when moose were more active across the landscape. From the fall questing period to the spring drop-off period, patches where colonization was high and extinction was low had higher proportions of young (shrub/forage) mixed forest at higher elevations. Further, we evaluated the fitness consequences of habitat selection by adult females during the fall questing period, when females and their calves acquire ticks. We compared Resource Selection Functions (RSF) for five females that successfully reared a calf to age 1 with five females whose calves perished due to ticks. Adult female moose whose offspring perished selected habitats in the fall that spatially coincided with areas of high occupancy probability during the spring tick drop-off period. In contrast, adult female moose whose offspring survived selected areas where the probability of occupancy during the spring drop-off was low; at present, natural selection may favor female adults who do not select the same habitats in fall as in spring. Our model coefficients and mapped results define “hotspots” that are likely encouraging the deleterious effects of the tick-moose cycle. These findings fill knowledge gaps about moose habitat selection that may improve the effectiveness of management aimed at reversing declining population trends.

Vermont

Grassification and fast-evolving fire connectivity and risk in the Sonoran Desert, United States

In the southwestern United States, non-native grass invasions have increased wildfire occurrence in deserts and the likelihood of fire spread to and from other biomes with disparate fire regimes. The elevational transition between desertscrub and montane grasslands, woodlands, and forests generally occurs at ∼1,200 masl and has experienced fast suburbanization and an expanding wildland-urban interface (WUI). In summer 2020, the Bighorn Fire in the Santa Catalina Mountains burned 486 km 2 and prompted alerts and evacuations along a 40-km stretch of WUI below 1,200 masl on the outskirts of Tucson, Arizona, a metropolitan area of >1M people. To better understand the changing nature of the WUI here and elsewhere in the region, we took a multidimensional and timely approach to assess fire dynamics along the Desertscrub-Semi-desert Grassland ecotone in the Catalina foothills, which is in various stages of non-native grass invasion. The Bighorn Fire was principally a forest fire driven by a long-history of fire suppression, accumulation of fine fuels following a wet winter and spring, and two decades of hotter droughts, culminating in the hottest and second driest summer in the 125-yr Tucson weather record. Saguaro ( Carnegia gigantea ), a giant columnar cactus, experienced high mortality. Resprouting by several desert shrub species may confer some post-fire resiliency in desertscrub. Buffelgrass and other non-native species played a minor role in carrying the fire due to the patchiness of infestation at the upper edge of the Desertscrub biome. Coupled state-and-transition fire-spread simulation models suggest a marked increase in both burned area and fire frequency if buffelgrass patches continue to expand and coalesce at the Desertscrub/Semi-desert Grassland interface. A survey of area residents six months after the fire showed awareness of buffelgrass was significantly higher among residents that were evacuated or lost recreation access, with higher awareness of fire risk, saguaro loss and declining property values, in that order. Sustained and timely efforts to document and assess fast-evolving fire connectivity due to grass invasions, and social awareness and perceptions, are needed to understand and motivate mitigation of an increasingly fire-prone future in the region.

Arizona

Flight altitudes of raptors in southern Africa highlight vulnerability of threatened species to wind turbines

Energy infrastructure, particularly for wind power, is rapidly expanding in Africa, creating the potential for conflict with at-risk wildlife populations. Raptor populations are especially susceptible to negative impacts of fatalities from wind energy because individuals tend to be long-lived and reproduce slowly. A major determinant of risk of collision between flying birds and wind turbines is the altitude above ground at which a bird flies. We examine 18,710 observations of flying raptors recorded in southern Africa and we evaluate, for 49 species, the frequency with which they were observed to fly at the general height of a wind turbine rotor-swept zone (50–150 m). Threatened species, especially vultures, were more likely to be observed at turbine height than were other species, suggesting that these raptors are most likely to be affected by wind power development across southern Africa. Our results highlight that threatened raptor species, particularly vultures, might be especially impacted by expanded wind energy infrastructure across southern Africa.

Frontiers in Ecology and Evolution

Citizen science data collection for integrated wildlife population analyses

Citizen science, or community science, has emerged as a cost-efficient method to collect data for wildlife monitoring. To inform research and conservation, citizen science sampling designs should collect data that match the robust statistical analyses needed to quantify species and population patterns. Further increasing the contributions of citizen science, integrating citizen science data with other datasets and datatypes can improve population estimates and expand the spatiotemporal extent of inference. We demonstrate these points with a citizen science program called iSeeMammals developed in New York state in 2017 to supplement costly systematic spatial capture-recapture sampling by collecting opportunistic data from one-off observations, hikes, and camera traps. iSeeMammals has initially focused on the growing population of American black bear ( Ursus americanus ), with integrated analysis of iSeeMammals camera trap data with systematic data for a region with a growing bear population. The triumvirate of increased spatial and temporal coverage by at least twofold compared to systematic sampling, an 83% reduction in annual sampling costs, and improved density estimates when integrated with systematic data highlight the benefits of collecting presence-absence data in citizen science programs for estimating population patterns. Additional opportunities will come from applying presence-only data, which are oftentimes more prevalent than presence-absence data, to integrated models. Patterns in data submission and filtering also emphasize the importance of iteratively evaluating patterns in engagement, usability, and accessibility, especially focusing on younger adult and teenage demographics, to improve data quality and quantity. We explore how the development and use of integrated models may be paired with citizen science project design in order to facilitate repeated use of datasets in standalone and integrated analyses for supporting wildlife monitoring and informing conservation.

Frontiers in Ecology and Evolution

Physiomorphic transformation in extreme endurance migrants: Revisiting the case of bar-tailed godwits preparing for trans-pacific flights

In a 1998 paper entitled “Guts don’t fly: small digestive organs in obese bar-tailed godwits,” Piersma and Gill (1998) showed that the digestive organs were tiny and the fat loads huge in individuals suspected of embarking on a non-stop flight from Alaska to New Zealand. It was suggested that prior to migratory departure, these godwits would shrink the digestive organs used during fuel deposition and boost the size and capacity of exercise organs to optimize flight performance. Here we document the verity of the proposed physiomorphic changes by comparing organ sizes and body composition of bar-tailed godwits Limosa lapponica baueri collected in modesty midway during their fueling period (mid-September; fueling, n = 7) with the previously published data for godwits that had just departed on their trans-Pacific flight (October 19; flying, n = 9). Mean total body masses for the two groups were nearly identical, but nearly half of the body mass of fueling godwits consisted of water, while fat constituted over half of total body mass of flying godwits. The two groups also differed in their fat-free mass components. The heart and flight muscles were heavier in fueling godwits, but these body components constituted a relatively greater fraction of the fat-free mass in flying godwits. In contrast, organs related to digestion and homeostasis were heavier in fueling godwits, and most of these organ groups were also relatively larger in fueling godwits compared to flying godwits. These results reflect the functional importance of organ and muscle groups related to energy acquisition in fueling godwits and the consequences of flight-related exertion in flying godwits. The extreme physiomorphic changes apparently occurred over a short time window (≤1 month). We conclude that the inferences made on the basis of the 1998 paper were correct. The cues and stimuli which moderate these changes remain to be studied.

Frontiers in Ecology and Evolution

Hierarchical functional response of a forager on a wetland landscape

We show that for some foragers the form that a functional response takes depends on the temporal and spatial scales considered. In representing the consumption rate of an organism, it may be necessary to use a hierarchy of functional responses. Consider, for example, a wading bird foraging in wetland landscape characterized by a spatial distribution of potential foraging sites, such as ponds. At the smallest time scale of minutes or hours, during which a wading bird is foraging within a single site, the functional response will reflect the local density of prey, as well as features of the site that affect the feeding rate, such as water depth. At this short time scale, which is determined by the giving up time of the wading bird in a particular site, prey density may be relatively constant. The food intake from a particular pond is then the product of the time spent before giving-up time and moving to another site and the rate of prey consumption at that site. A prey-centered functional response is most appropriate for describing the prey consumption rate. We propose that over the longer time scale of a day, during which a wading bird may visit several foraging sites, the type of functional response can be considered to be patch centered. That is, it is influenced by the spatial configuration of sites with available prey and the wading bird’s strategy of choosing among different sites and decisions on how long to stay in any given sites. Over the time scale of a day, if the prey densities stay relatively constant, the patch-centered functional response for a constant environment is adequate. However, on the longer time scale of a breeding season, in which changing water levels result in temporal changes in the availability of prey in sites, a third hierarchical level may be relevant. At that scale, the way in which the landscape pattern changes through time, and how the wading bird responds, influences the functional response. This hierarchical concept applies to a colony of breeding wading birds foraging in wetlands such as the Everglades.

Frontiers in Ecology and Evolution

Reconnecting the Elwha River: Spatial patterns of fish response to dam removal

The removal of two large dams on the Elwha River was completed in 2014 with a goal of restoring anadromous salmonid populations. Using observations from ongoing field studies, we compiled a timeline of migratory fish passage upstream of each dam. We also used spatially continuous snorkeling surveys in consecutive years before (2007, 2008) and after (2018, 2019) dam removal during summer baseflow to assess changes in fish distribution and density over 65 km of the mainstem Elwha River. Before dam removal, anadromous fishes were limited to the 7.9 km section of river downstream of Elwha Dam, potamodromous species could not migrate throughout the river system, and resident trout were the most abundant species. After dam removal, there was rapid passage into areas upstream of Elwha Dam, with 8 anadromous species (Chinook, Coho, Sockeye, Pink, Chum, Winter Steelhead, Summer Steelhead, Pacific Lamprey, and Bull Trout) observed within 2.5 years. All of these runs except Chum Salmon were also observed in upper Elwha upstream of Glines Canyon Dam within 5 years. The spatial extent of fish passage by adult Chinook Salmon and Summer Steelhead increased by 50 km and 60 km, respectively, after dam removal. Adult Chinook Salmon densities in some previously inaccessible reaches in the middle section of the river exceeded the highest densities observed in the lower section of the river prior to dam removal. The large number (>100) of adult Summer Steelhead in the upper river after dam removal was notable because it was among the rarest anadromous species in the Elwha River prior to dam removal. The spatial extent of trout and Bull Trout remained unchanged after dam removal, but their total abundance increased and their highest densities shifted from the lower 25 km of the river to the upper 40 km. Our results show that reconnecting the Elwha River through dam removal provided fish access to portions of the watershed that had been blocked for nearly a century.

Washington

Editorial: Environmental DNA innovations for conservation

Environmental DNA (eDNA) analysis refers to the collection of bulk environmental samples such as water, sediment, or air, and studying the genetic remnants that organisms have shed into their environment to gain information about species presence.

Frontiers in Ecology and Evolution

Antecedent climatic conditions spanning several years influence multiple land-surface phenology events in semi-arid environments

Ecological processes are complex, often exhibiting non-linear, interactive, or hierarchical relationships. Furthermore, models identifying drivers of phenology are constrained by uncertainty regarding predictors, interactions across scales, and legacy impacts of prior climate conditions. Nonetheless, measuring and modeling ecosystem processes such as phenology remains critical for management of ecological systems and the social systems they support. We used random forest models to assess which combination of climate, location, edaphic, vegetation composition, and disturbance variables best predict several phenological responses in three dominant land cover types in the U.S. Northwestern Great Plains (NWP). We derived phenological measures from the 25-year series of AVHRR satellite data and characterized climatic predictors (i.e., multiple moisture and/or temperature based variables) over seasonal and annual timeframes within the current year and up to 4 years prior. We found that antecedent conditions, from seasons to years before the current, were strongly associated with phenological measures, apparently mediating the responses of communities to current-year conditions. For example, at least one measure of antecedent-moisture availability [precipitation or vapor pressure deficit (VPD)] over multiple years was a key predictor of all productivity measures. Variables including longer-term lags or prior year sums, such as multi-year-cumulative moisture conditions of maximum VPD, were top predictors for start of season. Productivity measures were also associated with contextual variables such as soil characteristics and vegetation composition. Phenology is a key process that profoundly affects organism-environment relationships, spatio-temporal patterns in ecosystem structure and function, and other ecosystem dynamics. Phenology, however, is complex, and is mediated by lagged effects, interactions, and a diversity of potential drivers; nonetheless, the incorporation of antecedent conditions and contextual variables can improve models of phenology.

Idaho, Montana, Nebraska, North Dakota, South Dako

New indicators of ecological resilience and invasion resistance to support prioritization and management in the sagebrush biome, United States

Ecosystem transformations to altered or novel ecological states are accelerating across the globe. Indicators of ecological resilience to disturbance and resistance to invasion can aid in assessing risks and prioritizing areas for conservation and restoration. The sagebrush biome encompasses parts of 11 western states and is experiencing rapid transformations due to human population growth, invasive species, altered disturbance regimes, and climate change. We built on prior use of static soil moisture and temperature regimes to develop new, ecologically relevant and climate responsive indicators of both resilience and resistance. Our new indicators were based on climate and soil water availability variables derived from process-based ecohydrological models that allow predictions of future conditions. We asked: (1) Which variables best indicate resilience and resistance? (2) What are the relationships among the indicator variables and resilience and resistance categories? (3) How do patterns of resilience and resistance vary across the area? We assembled a large database ( n = 24,045) of vegetation sample plots from regional monitoring programs and derived multiple climate and soil water availability variables for each plot from ecohydrological simulations. We used USDA Natural Resources Conservation Service National Soils Survey Information, Ecological Site Descriptions, and expert knowledge to develop and assign ecological types and resilience and resistance categories to each plot. We used random forest models to derive a set of 19 climate and water availability variables that best predicted resilience and resistance categories. Our models had relatively high multiclass accuracy (80% for resilience; 75% for resistance). Top indicator variables for both resilience and resistance included mean temperature, coldest month temperature, climatic water deficit, and summer and driest month precipitation. Variable relationships and patterns differed among ecoregions but reflected environmental gradients; low resilience and resistance were indicated by warm and dry conditions with high climatic water deficits, and moderately high to high resilience and resistance were characterized by cooler and moister conditions with low climatic water deficits. The new, ecologically-relevant indicators provide information on the vulnerability of resources and likely success of management actions, and can be used to develop new approaches and tools for prioritizing areas for conservation and restoration actions.

Frontiers in Ecology and Evolution

A comprehensive multi-state conditional occupancy model for evaluating interactions of non-native and native species

A major challenge in ecology is disentangling interactions of non-native, potentially invasive species on native species. Conditional two-species occupancy models examine the effects of dominant species (e.g., non-native) on subordinate species (e.g., native) while considering the possibility that occupancy of one species may affect occupancy and/ or detection of the other. Although conditional two-species models are useful for evaluating the influence of one species on presence of another, it is possible that species interactions are density dependent. Therefore, we developed a novel two-species occupancy model that incorporates multiple abundance states (i.e., absent, present, abundant) of the native species. We showcase the utility of this model with a case study that incorporates random effects and covariates on both occupancy and detection to help disentangle species interactions given varying occupancy and detection in different abundance states. We use snorkel survey data from the Umpqua basin, Oregon, where it is hypothesized that smallmouth bass Micropterus dolomieu , a non-native piscivore, exclude Umpqua chub Oregonichthys kalawatseti , a small endemic minnow. From our two-species multi-state (2SMS) model, we concluded that average occupancy was low for both fishes, and that when non-native bass were present, overall native chub occupancy in the present (0.18 ± 0.05 SD) and abundant (0.19 ± 0.03) states was higher than when non-natives were absent (0.14 ± 0.02/ 0.08 ± 0.02), indicating the non-native was not excluding the native species. By incorporating a species interaction factor, we found a positive association (6.75 ± 5.54 SD) between native chub and non-native bass. The covariates strongly related to occupancy were elevation, algae, and land cover type (urban and shrub). Detection probability for both species (0.21–0.82) was most strongly related to the covariates day of year, water temperature, gravel substrate, and stream order/ magnitude. Incorporation of detection probability and covariates enabled interpretation of interactions between the two species that may have been missed without their inclusion in the modeling process. Our new 2SMS occupancy model can be used by scientists and managers with a broad range of survey and covariate data to disentangle species interactions problems to help them inform management decisions.

Frontiers in Ecology and Evolution