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Rapid microsatellite marker development using next generation pyrosequencing to inform invasive Burmese python -- Python molurus bivittatus -- management

Invasive species represent an increasing threat to native ecosystems, harming indigenous taxa through predation, habitat modification, cross-species hybridization and alteration of ecosystem processes. Additionally, high economic costs are associated with environmental damage, restoration and control measures. The Burmese python, Python molurus bivittatus, is one of the most notable invasive species in the US, due to the threat it poses to imperiled species and the Greater Everglades ecosystem. To address population structure and relatedness, next generation sequencing was used to rapidly produce species-specific microsatellite loci. The Roche 454 GS-FLX Titanium platform provided 6616 di-, tri- and tetra-nucleotide repeats in 117,516 sequences. Using stringent criteria, 24 of 26 selected tri- and tetra-nucleotide loci were polymerase chain reaction (PCR) amplified and 18 were polymorphic. An additional six cross-species loci were amplified, and the resulting 24 loci were incorporated into eight PCR multiplexes. Multi-locus genotypes yielded an average of 61% (39%–77%) heterozygosity and 3.7 (2–6) alleles per locus. Population-level studies using the developed microsatellites will track the invasion front and monitor population-suppression dynamics. Additionally, cross-species amplification was detected in the invasive Ball, P. regius, and Northern African python, P. sebae. These markers can be used to address the hybridization potential of Burmese pythons and the larger, more aggressive P. sebae.

Florida

Bioeconomic analysis of selected conservation practices on soil erosion and freshwater fisheries

Farmers can generate environmental benefits (improved water quality and fisheries and wildlife habitat), but they may not be able to quantify them. Furthermore, farmers may reduce their incomes from managing lands to produce these positive externalities but receive little monetary compensation in return. This study simulated the relationship between agricultural practices, water quality, fish responses to suspended sediment and farm income within two small watersheds, one of a cool water stream and one of a warm water stream. Using the Agricultural Drainage and Pesticide Transport (ADAPT) model, this study related best management practices (BMPs) to calculated instream suspended sediment concentrations by estimating sediment delivery, runoff, base flow, and streambank erosion to quantify the effects of suspended sediment exposure on fish communities. By implementing selected BMPs in each watershed, annual net farm income declined $18,000 to $28,000 (1 to 3 percent) from previous levels. “Lethal” fish events from suspended sediments in the cool water watershed decreased by 60 percent as conservation tillage and riparian buffers increased. Despite reducing suspended sediments by 25 percent, BMPs in the warm water watershed did not reduce the negative response of the fisheries. Differences in responses (physical and biological) between watersheds highlight potential gains in economic efficiency by targeting BMPs or by offering performance based “green payments.”

Journal of the American Water Resources Associatio

Characterization of fish assemblages in eleven multi-use reservoirs from North Carolina, USA

Managing impounded river systems is a recurring challenge for aquatic resource professionals because reservoirs serve multiple functions with different ecological and socioeconomic outcomes. However, research on fishes in reservoirs has disproportionally focused on recreationally and economically important species, with less attention directed toward fish assemblages despite the potential for management at the assemblage level. As such, evaluation of relationships between reservoir fish assemblages and biotic and abiotic factors and testing whether assemblage structure is affected by changing environmental conditions may deepen ecological understanding and provide insights for reservoir fisheries management. Our overall objective was to assess these relationships in 11 reservoirs from North Carolina, USA. We sampled fish assemblages in the reservoirs, which spanned five river basins representing a range of habitat conditions, using experimental gillnets and pulsed DC nighttime electrofishing. Multivariate statistical analyses indicated that taxonomic differences in fish assemblage composition among river basins followed a gradient of productivity. The top contributing species to reservoir dissimilarity were bluegill ( Lepomis macrochirus ), gizzard shad ( Dorosoma cepedianum ), black crappie ( Pomoxis nigromaculatus ), and white perch ( Morone americana ). These four species were positively associated with factors that reflect increasing eutrophic conditions in the 11 reservoirs and could, therefore, serve as indicators of reservoir productivity, anthropogenic influence, and fish assemblage structure, in addition to their key role in reservoir fisheries management. Whereas ­fisheries research has historically focused on assessing fish ­populations, our results illustrate the ecological and management insights derived from simultaneously collecting assemblage- and population-level data. Research on reservoir fish assemblages in relation to biotic and abiotic conditions may help advance fish ecology and management alike.

North Carolina

International river basins of the world

It is becoming acknowledged that water is likely to be the most pressing environmental concern of the next century. Difficulties in river basin management are only exacerbated when the resource crosses international boundaries. One critical aid in the assessment of international waters has been the Register of International Rivers a compendium which listed 214 international waterways that cover 47% of the earth's continental land surface. The Register, though, was last updated in 1978 by the now defunct United Nations Department of Economic and Social Affairs. The purpose of this paper is to update the Register in order to reflect the quantum changes that have taken place over the last 22 years, both in global geopolitics and in map coverage and technology. By accessing digital elevation models at spatial resolutions of 30 arc seconds, corroborating at a unified global map coverage of at least 1:1 000 000, and superimposing the results over complete coverage of current political boundaries, we are able to provide a new register which lists 261 international rivers, covering 45.3% of the land surface of the earth (excluding Antarctica). This paper lists all international rivers with their watershed areas, the nations which share each watershed,their respective territorial percentages, and notes on changes in or disputes over international boundaries since 1978.

International Journal of Water Resources Developme

Managing native and non-native sea lamprey ( Petromyzon marinus ) through anthropogenic change: A prospective assessment of key threats and uncertainties

Sea lamprey ( Petromyzon marinus ) is a species of conservation concern in their native range of the Atlantic coasts of Europe (Near Threatened to Critically Endangered) and North America (Secure to Critically Imperiled), and an invasive species of great economic and ecological concern in the Laurentian Great Lakes. Despite differences in life history strategy (anadromous natives vs adfluvial non-natives), the biology of sea lamprey is sufficiently similar to expect comparable responses to large-scale environmental change. We take a prospective look at the future (50 to 100 years) of sea lamprey management in an era of considerable environmental disturbance, and consider biological responses, management actions, and the future status of populations across the native and non-native ranges. Based on facilitated discussion by a diverse group of international experts, two major but poorly characterized classes of threats to sea lamprey were identified: climate change and socio-political issues. We discuss how climate induced changes affect growth, bioenergetics, and phenology of sea lamprey, and associated effects on control tactics (pesticides and barriers) and conservation. We consider tensions surrounding improving connectivity in the Great Lakes while controlling invasive sea lamprey, and discuss supplements and alternatives to pesticides and their wider effect, as well as the effects of new invasive species. To prevent the extirpation of native sea lamprey populations, or the re-expansion of non-native populations, we conclude with a call for new and ongoing dialogue and collaboration among all sea lamprey biologists and managers across the native and non-native range.

Journal of Great Lakes Research

Consequences of changing water clarity on the fish and fisheries of the Laurentian Great Lakes

Human-driven environmental change underlies recent changes in water clarity in many of the world’s great lakes, yet our understanding of the consequences of these changes on the fish and fisheries they support remains incomplete. Herein, we offer a framework to organize current knowledge, guide future research, and help fisheries managers understand how water clarity can affect their valued populations. Emphasizing Laurentian Great Lakes findings where possible, we describe how changing water clarity can directly affect fish populations and communities by altering exposure to ultraviolet radiation, foraging success, predation risk, reproductive behavior, or territoriality. We also discuss how changing water clarity can affect fisheries harvest and assessment through effects on fisher behavior and sampling efficiency (i.e., catchability). Finally, we discuss whether changing water clarity can affect understudied aspects of fishery performance, including economic and community benefits. We conclude by identifying generalized predictions and discuss their implications for priority research questions for the Laurentian Great Lakes. Even though the motivation for this work was regional, the breadth of the review and generality of the framework are readily transferable to other freshwater and marine habitats.

Great Lakes

Harvesting and replenishment policies for renewable natural resources

The current paper links the optimal intertemporal use of renewable natural resources to the harvesting activities of various economic agents. Previous contributions cite market forces as a causative factor inducing the extirpation of renewable natural resources. The analysis given here discusses investment in the stock of renewable resources and cites important examples of this activity. By introducing joint harvesting and replenishment strategies into a model of renewable resource use, the analysis adds descriptive reality and relevance to positive and normative discussions of renewable natural resource use. A high price for the yield or a high discount rate tend to diminish the size of the optimum stationary stock of the resource with a non-replenishment harvesting strategy. Optimal non-replenishment harvesting strategies for renewable natural resources will exhaustion or extirpation of the resource if the price of the yield or the discount rate are sufficiently large. However, the availability of a replenishment technology and the use of replenishment activities tends to buffer the resource against exhaustion or extirpation.

Journal of Environmental Management

Valuation of the flood attenuation ecosystem service in Difficult Run, VA, USA

Floodplains and riparian wetlands provide several ecosystem services that directly benefit people. We present a methodology for valuing the flood attenuation ecosystem service in Difficult Run, a suburban watershed with extensive natural floodplains in northern Virginia. High-resolution lidar-derived data were combined with GIS modeling techniques to produce estimates of flood inundation. We combined the modeled estimates with parcel-level property and primary economic data using a baseline and a counterfactual scenario to estimate the magnitude of flood attenuation and the associated value of the ecosystem service. Our framework brings new models and data to look at floodplains and an alternative land surface scenario in a way that has not previously been done. Annualized avoided property losses totaled $42,184 in the baseline scenario and $115,596 in the counterfactual scenario for the combined 200-, 100-, 50-, 20-, 10-, and 5-year flood events. We estimate the total annualized value of the flood attenuation ecosystem service in Difficult Run is $73,412, which is $77 per hectare of floodplain area and is consistent with similar valuation studies of floodplains. The framework presented here is not specific to the study area and could be deployed at larger spatial areas in other locations. Our methods may better inform land use decision making on the impacts of development in and surrounding floodplain areas.

Virginia

Grass carp (Ctenopharyngodon idella) movement states and backwater use can inform removal efforts

Expansion of grass carp ( Ctenopharyngodon idella ) within the Laurentian Great Lakes poses significant ecological and economic risks. Suppression efforts to remove grass carp have increased to curtail population growth, but capture efficiency remains low. One management challenge is the lack of understanding how this invasive species is using its novel habitats in the Great Lakes. This study aimed to identify spatiotemporal patterns in grass carp behavior and backwater use that could be exploited by management efforts, including removal. We used acoustic telemetry to examine when grass carp occurred in areas vulnerable to removal by boat electrofishing in the Sandusky River, Ohio, USA. Hidden Markov models were used to categorize behavioral states and evaluate their relationships with environmental factors. Best performing 2-state and 3-state hidden Markov models included proximity to backwater as a factor in state-dependent step length and transition probability between states, with a higher probability of being in less mobile states during the middle of the day and in backwaters. For the 2-state model, this less mobile state was also more common during warmer temperatures. Distinct diel patterns and backwater use identified in this study provide critical insights for improving management. Model results suggest that current removal efforts, focused primarily on the mainstem of the river, may benefit from adjustments in both timing and removal gear. The findings raise further questions about unobserved behaviors contributing to state assignment and sufficiently flexible adaptive management strategies to account for individual variation in grass carp behavior.

Ohio

The rise and fall of water hyacinth in Lake Victoria and the Kagera River basin, 1989-2001

Water hyacinth (Eichhornia crassipes (Mart.) Solms) is an invasive aquatic macrophyte associated with major negative economic and ecological impacts to the Lake Victoria region since the plant's establishment in Uganda in the 1980s. Reliable estimates of water hyacinth distribution and extent are required to gauge the severity of the problem through time, relate water hyacinth abundance to environmental factors, identify areas requiring management action, and assess the efficacy of management actions. To provide such estimates and demonstrate the utility of remote sensing for this application, we processed and analyzed remotely sensed imagery to determine the distribution and extent of water hyacinth. Maps were produced and coverage was quantified using a hybrid unsupervised image classification approach with manual editing for each of the riparian countries of Kenya, Tanzania, and Uganda, as well as for numerous gulfs and bays. A similar procedure was carried out for selected lakes in the Rwanda-Tanzania borderlands lakes region in the Kagera River basin. Results confirm the severity of the water hyacinth infestation, especially in the northern parts of the lake. A maximum lake-wide extent of at least 17,374 ha was attained in 1998. Following this, a combination of factors, including conditions associated with the 1997 to 1998 El Nin??o and biocontrol with water hyacinth weevils, appear to have contributed to a major decline in water hyacinth in the most affected parts of the lake. Some lakes in the Kagera basin, such as Lake Mihindi, Rwanda, were severely infested in the late 1990s, but the level of infestation in most of these decreased markedly by the early 2000s.

Journal of Aquatic Plant Management

Coupling ecological and social network models to assess “transmission” and “contagion” of an aquatic invasive species

Network analysis is used to address diverse ecological, social, economic, and epidemiological questions, but few efforts have been made to combine these field-specific analyses into interdisciplinary approaches that effectively address how complex systems are interdependent and connected to one another. Identifying and understanding these cross-boundary connections improves natural resource management and promotes proactive, rather than reactive, decisions. This research had two main objectives; first, adapt the framework and approach of infectious disease network modeling so that it may be applied to the socio-ecological problem of spreading aquatic invasive species, and second, use this new coupled model to simulate the spread of the invasive Chinese mystery snail ( Bellamya chinensis ) in a reservoir network in Southeastern Nebraska, USA. The coupled model integrates an existing social network model of how anglers move on the landscape with new reservoir-specific ecological network models. This approach allowed us to identify 1) how angler movement among reservoirs aids in the spread of B . chinensis , 2) how B . chinensis alters energy flows within individual-reservoir food webs, and 3) a new method for assessing the spread of any number of non-native or invasive species within complex, social-ecological systems.

Journal of Environmental Management

Biological invasions, ecological resilience and adaptive governance

In a world of increasing interconnections in global trade as well as rapid change in climate and land cover, the accelerating introduction and spread of invasive species is a critical concern due to associated negative social and ecological impacts, both real and perceived. Much of the societal response to invasive species to date has been associated with negative economic consequences of invasions. This response has shaped a war-like approach to addressing invasions, one with an agenda of eradications and intense ecological restoration efforts towards prior or more desirable ecological regimes. This trajectory often ignores the concept of ecological resilience and associated approaches of resilience-based governance. We argue that the relationship between ecological resilience and invasive species has been understudied to the detriment of attempts to govern invasions, and that most management actions fail, primarily because they do not incorporate adaptive, learning-based approaches. Invasive species can decrease resilience by reducing the biodiversity that underpins ecological functions and processes, making ecosystems more prone to regime shifts. However, invasions do not always result in a shift to an alternative regime; invasions can also increase resilience by introducing novelty, replacing lost ecological functions or adding redundancy that strengthens already existing structures and processes in an ecosystem. This paper examines the potential impacts of species invasions on the resilience of ecosystems and suggests that resilience-based approaches can inform policy by linking the governance of biological invasions to the negotiation of tradeoffs between ecosystem services.

Journal of Environmental Management

Spatially explicit modeling of blackbird abundance in the Prairie Pothole Region

Knowledge of factors influencing animal abundance is important to wildlife biologists developing management plans. This is especially true for economically important species such as blackbirds (Icteridae), which cause more than $100 million in crop damages annually in the United States. Using data from the North American Breeding Bird Survey, the National Land Cover Dataset, and the National Climatic Data Center, we modeled effects of regional environmental variables on relative abundance of 3 blackbird species (red-winged blackbird, Agelaius phoeniceus ; yellow-headed blackbird, Xanthocephalus xanthocephalus ; common grackle, Quiscalus quiscula ) in the Prairie Pothole Region of the central United States. We evaluated landscape covariates at 3 logarithmically related spatial scales (1,000 ha, 10,000 ha, and 100,000 ha) and modeled weather variables at the 100,000-ha scale. We constructed models a priori using information from published habitat associations. We fit models with WinBUGS using Markov chain Monte Carlo techniques. Both landscape and weather variables contributed strongly to predicting blackbird relative abundance (95% credibility interval did not overlap 0). Variables with the strongest associations with blackbird relative abundance were the percentage of wetland area and precipitation amount from the year before bird surveys were conducted. The influence of spatial scale appeared small—models with the same variables expressed at different scales were often in the best model subset. This large-scale study elucidated regional effects of weather and landscape variables, suggesting that management strategies aimed at reducing damages caused by these species should consider the broader landscape, including weather effects, because such factors may outweigh the influence of localized conditions or site-specific management actions. The regional species distributional models we developed for blackbirds provide a tool for understanding these broader landscape effects and guiding wildlife management practices to areas that are optimally beneficial. Published 2015. This article is a U.S. Government work and is in the public domain in the USA.

Iowa, Minnesotta, Montana, Nebraska, North Dakota

A comparative and experimental evaluation of performance of stocked diploid and triploid brook trout

Despite numerous negative impacts, nonnative trout are still being stocked to provide economically and socially valuable sport fisheries in western mountain lakes. We evaluated relative performance and potential differences in feeding strategy and competitive ability of triploid versus diploid brook trout Salvelinus fontinalis in alpine lakes, as well as behavioral and performance differences of diploid and triploid brook trout in two controlled experimental settings: behavioral experiments in the laboratory and performance evaluations in ponds. Across lakes, catch per unit effort (CPUE) and relative weight (Wr ) were not significantly different between ploidy levels. Mean sizes were also similar between ploidy levels except in two of the larger lakes where diploids attained slightly larger sizes (approximately 20 mm longer). We observed no significant differences between diploids and triploids in diet, diet preference, or trophic structure. Similarly, growth and condition did not differ between ploidy levels in smaller-scale pond experiments, and aggressive behavior did not differ between ploidy levels (fed or unfed fish trials) in the laboratory. Independent of ploidy level, the relative performance of brook trout varied widely among lakes, a pattern that appeared to be a function of lake size or a factor that covaries with lake size such as temperature regime or carrying capacity. In summary, we observed no significant differences in the relative performance of brook trout from either ploidy level across a number of indices, systems, and environmental conditions, nor any indication that one group is more aggressive or a superior competitor than the other. Collectively, these results suggest that triploid brook trout will offer a more risk-averse and promising management opportunity when they are stocked to these lakes and elsewhere to simultaneously meet the needs for the sport fishery and conservation objectives.

North American Journal of Fisheries Management

Antibiotic resistant bacteria in wildlife: Perspectives on trends, acquisitions and dissemination, data gaps, and future directions

The proliferation of antibiotic resistant bacteria in the environment has potential negative economic and health consequences. Thus, previous investigations have targeted wild animals to understand the occurrence of antibiotic resistance in diverse environmental sources. In this critical review and synthesis, we summarize important concepts learned through the sampling of wildlife for antibiotic resistant indicator bacteria. These concepts are helpful for understanding dissemination of resistance through environmental pathways and helping to guide future research efforts. Our review is comprised of six sections. The first section briefly introduces antibiotic resistance as it pertains to bacteria harbored in environmental sources such as wild animals. Next, we differentiate wildlife from other animals in the context of how diverse taxa provide different information on antibiotic resistance in the environment. In the third section, we identify representative research and seminal works that illustrate important associations between the occurrence of antibiotic resistant bacteria in wildlife and anthropogenic inputs into the environment. For example, we highlight numerous investigations that support the premise that anthropogenic inputs into the environment drive the occurrence of antibiotic resistance in bacteria harbored by free-ranging wildlife. Additionally, we summarize previous research demonstrating foraging as a mechanism by which wildlife may be exposed to anthropogenic antibiotic resistance contamination in the environment. In the fourth section of our review, we summarize molecular evidence for the acquisition and dissemination of resistance among bacteria harbored by wildlife. In the fifth section, we identify what we believe to be important data gaps and potential future directions that other researchers may find useful towards the development of efficient, informative, and impactful investigations of antibiotic resistant bacteria in wildlife. Finally, we conclude our review by highlighting the need to move from surveys that simply identify antibiotic resistant bacteria in wildlife towards hypothesis-driven investigations that: (1) identify point sources of antibiotic resistance; (2) provide information on risk to human and animal health; (3) identify interventions that may interrupt environmentally mediated pathways of antibiotic resistance acquisition/transmission; and (4) evaluate whether management practices are leading to desirable outcomes.

Journal of Wildlife Diseases

A method to quantify and value floodplain sediment and nutrient retention ecosystem services

Floodplains provide critical ecosystem services to local and downstream communities by retaining floodwaters, sediments, and nutrients. The dynamic nature of floodplains is such that these areas can both accumulate sediment and nutrients through deposition, and export material downstream through erosion. Therefore, estimating floodplain sediment and nutrient retention should consider the net flux of both depositional and erosive processes. An ecosystem services framework was used to quantify and value the sediment and nutrient ecosystem service provided by floodplains in the Difficult Run watershed, a small (151 km 2 ) suburban watershed located in the Piedmont of Virginia (USA). A sediment balance was developed for Difficult Run and two nested watersheds. The balance included upland sediment delivery to streams, stream bank flux, floodplain flux, and stream load. Upland sediment delivery was estimated using geospatial datasets and a modified Revised Universal Soil Loss Equation. Predictive models were developed to extrapolate field measurements of the flux of sediment, sediment-bound nitrogen (N), and sediment-bound phosphorus (P) from stream banks and floodplains to 3232 delineated stream segments in the study area. A replacement cost approach was used to estimate the economic value of the sediment and nutrient retention ecosystem service based on estimated net stream bank and floodplain flux of sediment-bound N for all streams in the study area. Results indicated the net fluvial fluxes of sediment, sediment-bound N, and sediment-bound P were −10,439 Mg yr −1 (net export), 57,300 kg-N yr −1 (net trapping), and 98 kg-P yr −1 (net trapping), respectively. For sediment, floodplain retention was offset by substantial losses from stream bank erosion, particularly in headwater catchments, resulting in a net export of sediment. Nutrient retention in the floodplain exceeded that lost through stream bank erosion resulting in net retention of nutrients (TN and TP). Using a conservative cost estimate of \$12.69 (USD) per kilogram of nitrogen, derived from wastewater treatment costs, the estimated annual value for sediment and nutrient retention on Difficult Run floodplains was \$727,226 ± 194,220 USD/yr. Values and differences in floodplain nitrogen retention among stream reaches can be used to target areas for floodplain conservation and stream restoration. The methods presented are scalable and transferable to other areas if appropriate datasets are available for validation.

Virginia

Harvesting wildlife affected by climate change: a modelling and management approach for polar bears

The conservation of many wildlife species requires understanding the demographic effects of climate change, including interactions between climate change and harvest, which can provide cultural, nutritional or economic value to humans. We present a demographic model that is based on the polar bear Ursus maritimus life cycle and includes density-dependent relationships linking vital rates to environmental carrying capacity ( K ). Using this model, we develop a state-dependent management framework to calculate a harvest level that (i) maintains a population above its maximum net productivity level (MNPL; the population size that produces the greatest net increment in abundance) relative to a changing K , and (ii) has a limited negative effect on population persistence. Our density-dependent relationships suggest that MNPL for polar bears occurs at approximately 0·69 (95% CI = 0·63–0·74) of K . Population growth rate at MNPL was approximately 0·82 (95% CI = 0·79–0·84) of the maximum intrinsic growth rate, suggesting relatively strong compensation for human-caused mortality. Our findings indicate that it is possible to minimize the demographic risks of harvest under climate change, including the risk that harvest will accelerate population declines driven by loss of the polar bear's sea-ice habitat. This requires that (i) the harvest rate – which could be 0 in some situations – accounts for a population's intrinsic growth rate, (ii) the harvest rate accounts for the quality of population data (e.g. lower harvest when uncertainty is large), and (iii) the harvest level is obtained by multiplying the harvest rate by an updated estimate of population size. Environmental variability, the sex and age of removed animals and risk tolerance can also affect the harvest rate. Synthesis and applications . We present a coupled modelling and management approach for wildlife that accounts for climate change and can be used to balance trade-offs among multiple conservation goals. In our example application to polar bears experiencing sea-ice loss, the goals are to maintain population viability while providing continued opportunities for subsistence harvest. Our approach may be relevant to other species for which near-term management is focused on human factors that directly influence population dynamics within the broader context of climate-induced habitat degradation.

Journal of Applied Ecology

Factors influencing elk recruitment across ecotypes in the Western United States

Ungulates are key components in ecosystems and economically important for sport and subsistence harvest. Yet the relative importance of the effects of weather conditions, forage productivity, and carnivores on ungulates are not well understood. We examined changes in elk ( Cervus canadensis ) recruitment (indexed as age ratios) across 7 states and 3 ecotypes in the northwestern United States during 1989–2010, while considering the effects of predator richness, forage productivity, and precipitation. We found a broad‐scale, long‐term decrease in elk recruitment of 0.48 juveniles/100 adult females/year. Weather conditions (indexed as summer and winter precipitation) showed small, but measurable, influences on recruitment. Forage productivity on summer and winter ranges (indexed by normalized difference vegetation index [NDVI] metrics) had the strongest effect on elk recruitment relative to other factors. Relationships between forage productivity and recruitment varied seasonally and regionally. The productivity of winter habitat was more important in southern parts of the study area, whereas annual variation in productivity of summer habitat had more influence on recruitment in northern areas. Elk recruitment varied by up to 15 juveniles/100 adult females across the range of variation in forage productivity. Areas with more species of large carnivores had relatively low elk recruitment, presumably because of increased predation. Wolves ( Canis lupus ) were associated with a decrease of 5 juveniles/100 adult females, whereas grizzly bears ( Ursus arctos ) were associated with an additional decrease of 7 juveniles/100 adult females. Carnivore species can have a critical influence on ungulate recruitment because their influence rivals large ranges of variation in environmental conditions. A more pressing concern, however, stems from persistent broad‐scale decreases in recruitment across the distribution of elk in the northwestern United States, irrespective of carnivore richness. Our results suggest that wildlife managers interested in improving recruitment of elk consider the combined effects of habitat and predators. Efforts to manage summer and winter ranges to increase forage productivity may have a positive effect on recruitment.

Journal of Wildlife Management