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Spatial occupancy models for predicting metapopulation dynamics and viability following reintroduction

The reintroduction of a species into its historic range is a critical component of conservation programmes designed to restore extirpated metapopulations. However, many reintroduction efforts fail, and the lack of rigorous monitoring programmes and statistical models have prevented a general understanding of the factors affecting metapopulation viability following reintroduction. Spatially explicit metapopulation theory provides the basis for understanding the dynamics of fragmented populations linked by dispersal, but the theory has rarely been used to guide reintroduction programmes because most spatial metapopulation models require presence&ndash;absence data from every site in the network, and they do not allow for observation error such as imperfect detection. We develop a spatial occupancy model that relaxes these restrictive assumptions and allows for inference about metapopulation extinction risk and connectivity. We demonstrate the utility of the model using six years of data on the Chiricahua leopard frog Lithobates chiricahuensis , a threatened desert-breeding amphibian that was reintroduced to a network of sites in Arizona USA in 2003. Our results indicate that the model can generate precise predictions of extinction risk and produce connectivity maps that can guide conservation efforts following reintroduction. In the case of L. chiricahuensis , many sites were functionally isolated, and 82% of sites were characterized by intermittent water availability and high local extinction probabilities (0&middot;84, 95% CI: 0&middot;64&ndash;0&middot;99). However, under the current hydrological conditions and spatial arrangement of sites, the risk of metapopulation extinction is estimated to be <3% over a 50-year time horizon. Low metapopulation extinction risk appears to result from the high dispersal capability of the species, the high density of sites in the region and the existence of predator-free permanent wetlands with low local extinction probabilities. Should management be required, extinction risk can be reduced by either increasing the hydroperiod of existing sites or by creating new sites to increase connectivity. Synthesis and applications . This work demonstrates how spatio-temporal statistical models based on ecological theory can be applied to forecast the outcomes of conservation actions such as reintroduction. Our spatial occupancy model should be particularly useful when management agencies lack the funds to collect intensive individual-level data.

Arizona, New Mexico

Applying a Bayesian weighted surveillance approach to detect chronic wasting disease in white‐tailed deer

Surveillance is critical for early detection of emerging and re‐emerging infectious diseases. Weighted surveillance leverages heterogeneity in infection risk to increase sampling efficiency. Here, we apply a Bayesian approach to estimate weights for 16 surveillance classes of white‐tailed deer in Wisconsin, USA, relative to hunter‐harvested yearling males. We used these weights to conduct a surveillance programme for detecting chronic wasting disease (CWD) in white‐tailed deer at Shenandoah National Park (SHEN) in Virginia, USA. Generally, for surveillance, risk of infection increased with age and was greater in males. Clinical suspect deer had the highest risk, with weight estimates of 33.33 and 9.09 for community‐reported and hunter‐reported suspect deer, respectively. Fawns had the lowest risk with an estimated weight of 0.001. We used surveillance weights for Wisconsin deer to determine sampling effort required to detect a CWD‐positive case in SHEN if prevalence in yearling males ≥0.025. The sampling required to detect CWD was 37–91 adult deer, depending on the adult male:female ratio in the surveillance stream. We collected rectal biopsies from 49 female and 21 male adult deer, and 10 additional samples from vehicle‐killed deer. CWD was not detected and we concluded with 95% probability that prevalence in the reference population (yearling males) was between 0.0% and 3.6%. Synthesis and applications . Our approach allows managers to estimate relative surveillance weights for different host classes and quantify limits of disease detection in real time when only a sample of animals from a population can be tested, resulting in considerable cost savings for agencies performing wildlife disease detection surveillance. Additionally, it provides a rigorous means of estimating prevalence limits when a disease/pathogen is not detected in a sample set. It is therefore applicable to other wildlife, domestic animal and human disease systems, which can be characterized by surveillance classes with heterogeneous probability of infection. This methodology is also extendable to other disciplines such as invasive species, environmental toxicology, and generally, any ecological question seeking to efficiently use scarce financial and human resources to maximize the detection probability of a rare event.

Journal of Applied Ecology

Invasive species management restores a plant-pollinator mutualism in Hawaii

1.The management and removal of invasive species may give rise to unanticipated changes in plant–pollinator mutualisms because they can alter the composition and functioning of plant–pollinator interactions in a variety of ways. To utilize a functional approach for invasive species management, we examined the restoration of plant–pollinator mutualisms following the large-scale removal of an invasive nectar thief and arthropod predator, Vespula pensylvanica. 2.We reduced V. pensylvanica populations in large plots managed over multiple years to examine the response of plant–pollinator mutualisms and the fruit production of a functionally important endemic Hawaiian tree species, Metrosideros polymorpha. To integrate knowledge of the invader's behaviour and the plant's mating system, we determined the efficacy of V. pensylvanica as a pollinator of M. polymorpha and quantified the dependence of M. polymorpha on animal pollination (e.g. level of self-compatibility and pollen limitation). 3.The reduction of V. pensylvanica in managed sites, when compared to unmanaged sites, resulted in a significant increase in the visitation rates of effective bee pollinators (e.g. introduced Apis mellifera and native Hylaeus spp.) and in the fruit production of M. polymorpha. 4.Apis mellifera, following the management of V. pensylvanica, appears to be acting as a substitute pollinator for M. polymorpha, replacing extinct or threatened bird and bee species in our study system. 5.Synthesis and applications. Fruit production of the native M. polymorpha was increased after management of the invasive pollinator predator V. pensylvanica; however, the main pollinators were no longer native but introduced. This research thus demonstrates the diverse impacts of introduced species on ecological function and the ambiguous role they play in restoration. We recommend incorporating ecological function and context into invasive species management as this approach may enable conservation managers to simultaneously minimize the negative and maximize the positive impacts (e.g. taxon substitution) of introduced species. Such novel restoration approaches are needed, especially in highly degraded ecosystems.

Hawai'i

How much is new information worth? Evaluating the financial benefit of resolving management uncertainty

Conservation decision-makers face a trade-off between spending limited funds on direct management action, or gaining new information in an attempt to improve management performance in the future. Value-of-information analysis can help to resolve this trade-off by evaluating how much management performance could improve if new information was gained. Value-of-information analysis has been used extensively in other disciplines, but there are only a few examples where it has informed conservation planning, none of which have used it to evaluate the financial value of gaining new information. We address this gap by applying value-of-information analysis to the management of a declining koala Phascolarctos cinereus population. Decision-makers responsible for managing this population face uncertainty about survival and fecundity rates, and how habitat cover affects mortality threats. The value of gaining new information about these uncertainties was calculated using a deterministic matrix model of the koala population to find the expected population growth rate if koala mortality threats were optimally managed under alternative model hypotheses, which represented the uncertainties faced by koala managers. Gaining new information about survival and fecundity rates and the effect of habitat cover on mortality threats will do little to improve koala management. Across a range of management budgets, no more than 1&middot;7% of the budget should be spent on resolving these uncertainties. The value of information was low because optimal management decisions were not sensitive to the uncertainties we considered. Decisions were instead driven by a substantial difference in the cost efficiency of management actions. The value of information was up to forty times higher when the cost efficiencies of different koala management actions were similar. Synthesis and applications . This study evaluates the ecological and financial benefits of gaining new information to inform a conservation problem. We also theoretically demonstrate that the value of reducing uncertainty is highest when it is not clear which management action is the most cost efficient. This study will help expand the use of value-of-information analyses in conservation by providing a cost efficiency metric by which to evaluate research or monitoring.

Journal of Applied Ecology

Ecological feedbacks can reduce population-level efficacy of wildlife fertility control

1. Anthropogenic stress on natural systems, particularly the fragmentation of landscapes and the extirpation of predators from food webs, has intensified the need to regulate abundance of wildlife populations with management. Controlling population growth using fertility control has been considered for almost four decades, but nearly all research has focused on understanding effects of fertility control agents on individual animals. Questions about the efficacy of fertility control as a way to control populations remain largely unanswered. 2. Collateral consequences of contraception can produce unexpected changes in birth rates, survival, immigration and emigration that may reduce the effectiveness of regulating animal abundance. The magnitude and frequency of such effects vary with species-specific social and reproductive systems, as well as connectivity of populations. Developing models that incorporate static demographic parameters from populations not controlled by contraception may bias predictions of fertility control efficacy. 3. Many population-level studies demonstrate that changes in survival and immigration induced by fertility control can compensate for the reduction in births caused by contraception. The most successful cases of regulating populations using fertility control come from applications of contraceptives to small, closed populations of gregarious and easily accessed species. 4. Fertility control can result in artificial selection pressures on the population and may lead to long-term unintentional genetic consequences. The magnitude of such selection is dependent on individual heritability and behavioural traits, as well as environmental variation. 5. Synthesis and applications . Understanding species' life-history strategies, biology, behavioural ecology and ecological context is critical to developing realistic expectations of regulating populations using fertility control. Before time, effort and funding are invested in wildlife contraception, managers may need to consider the possibility that many species and populations can compensate for reduction in fecundity, and this could minimize any reduction in population growth rate.

Journal of Applied Ecology

Synchrony — An emergent property of recreational fisheries

Recreational fisheries are traditionally managed at local scales, but more effective management could be achieved using a cross‐scale approach. To do this, we must first understand how local processes scale up to influence landscape patterns between anglers and resources. We highlight how population‐based synchrony methods, used in conjunction with a complex‐adaptive‐systems framework, can reveal emergent spatial properties within social‐ecological systems such as recreational fisheries. Herein, we quantified the level of spatial synchrony in angler behaviour, defined the relationship between angler synchrony and distance among waterbodies, and highlighted social‐ecological attributes contributing to these patterns. We leveraged a 111 waterbody‐year (34 waterbodies, 5‐year collection period) recreational fisheries dataset from Nebraska, USA to address these objectives. Intra‐annual patterns in angler behaviour were moderately synchronous across large spatial scales and predominately unrelated to distance among waterbodies. Large‐scale synchronous patterns in angler behaviour emerged from local‐scale interactions between angler heterogeneity and waterbody diversity. Spatial synchrony in angler behaviour is an emergent property that resulted from local‐level processes that scaled up to form large‐scale patterns. We posit that angler utility in combination with waterbodies sharing these desired utility components caused spatial synchrony among anglers with similar preferences or specializations. The level of spatial synchrony in angler behaviour will therefore depend on the degree of angler heterogeneity and waterbody diversity on the landscape, with high or low levels of both leading to low and high levels of spatial synchrony respectively. Synthesis and applications. Synchrony‐based methods proved useful for unveiling an emergent property in recreational fisheries that is beneficial for effective cross‐scale management. It may not be appropriate to extrapolate information and apply uniform management actions among local waterbodies because angler behaviour was not synchronous at small scales. Rather, anglers respond uniquely to waterbody diversity and therefore substitute waterbodies may be dispersed throughout the landscape. Creating boat access, for example could yield unintended consequences for a particular angler group and cause local and regional shifts in angler behaviour. Evaluating appropriate management options will require a cross‐scale monitoring approach that captures angler heterogeneity and waterbody diversity at multiple scales. Recreational fisheries are traditionally managed at local scales, but more effective management could be achieved using a cross‐scale approach. To do this, we must first understand how local processes scale up to influence landscape patterns between anglers and resources. We highlight how population‐based synchrony methods, used in conjunction with a complex‐adaptive‐systems framework, can reveal emergent spatial properties within social‐ecological systems such as recreational fisheries. Herein, we quantified the level of spatial synchrony in angler behaviour, defined the relationship between angler synchrony and distance among waterbodies, and highlighted social‐ecological attributes contributing to these patterns. We leveraged a 111 waterbody‐year (34 waterbodies, 5‐year collection period) recreational fisheries dataset from Nebraska, USA to address these objectives. Intra‐annual patterns in angler behaviour were moderately synchronous across large spatial scales and predominately unrelated to distance among waterbodies. Large‐scale synchronous patterns in angler behaviour emerged from local‐scale interactions between angler heterogeneity and waterbody diversity. Spatial synchrony in angler behaviour is an emergent property that resulted from local‐level processes that scaled up to form large‐scale patterns. We posit that angler

Journal of Applied Ecology

Negative frequency-dependent foraging behaviour in a generalist herbivore (Alces alces) and its stabilizing influence on food-web dynamics

1. Resource selection is widely appreciated to be context‐dependent and shaped by both biological and abiotic factors. However, few studies have empirically assessed the extent to which selective foraging behaviour is dynamic and varies in response to environmental conditions for free‐ranging animal populations. 2. Here, we assessed the extent that forage selection fluctuated in response to different environmental conditions for a free‐ranging herbivore, moose (Alces alces), in Isle Royale National Park, over a 10‐year period. More precisely, we assessed how moose selection for coniferous versus deciduous forage in winter varied between geographic regions and in relation to (a) the relative frequency of forage types in the environment (e.g. frequency‐dependent foraging behaviour), (b) moose abundance, (c) predation rate (by grey wolves) and (d) snow depth. These factors are potentially important for their influence on the energetics of foraging. We also built a series of food‐chain models to assess the influence of dynamic foraging strategies on the stability of food webs. 3. Our analysis indicates that moose exhibited negative frequency dependence, by selectively exploiting rare resources. Frequency‐dependent foraging was further mediated by density‐dependent processes, which are likely to be predation, moose abundance or some combination of both. In particular, frequency dependence was weaker in years when predation risk was high (i.e. when the ratio of moose to wolves was relatively low). Selection for conifers was also slightly weaker during deep snow years. 4. The food‐chain analysis indicates that the type of frequency‐dependent foraging strategy exhibited by herbivores had important consequences for the stability of ecological communities. In particular, the dynamic foraging strategy that we observed in the empirical analysis (i.e. negative frequency dependence being mediated by density‐dependent processes) was associated with more stable food web dynamics compared to fixed foraging strategies. 5. The results of this study indicated that forage selection is a complex ecological process, varying in response to both biological (predation and moose density) and abiotic factors (snow depth) and over relatively small spatial scales (between regions). This study also provides a useful framework for assessing the influence of other aspects of foraging behaviour on the stability of food web dynamics.

Michigan

Delineating ecologically-distinct groups for annual cycle management of a declining shorebird

1. Patterns of migratory connectivity are increasingly used to understand and manage threats throughout the annual cycle of migratory species. Strong migratory connectivity refers to when individuals from different populations remain spatially separated across the annual cycle, which may expose populations to unique sets of threats and conditions that cause differential population trends. However, the populations or groups used for species’ management are often defined a priori based on expert knowledge and/or management units, which may mask important population segregation and obscure differential population trends and their drivers. 2. We compared three approaches to defining management groups of a declining shorebird, the long-billed curlew ( Numenius americanus ), for annual cycle management: by expert-opinion, according to management flyways, and with unsupervised clustering of satellite tracking data that maximizes the strength of migratory connectivity. 3. Despite the curlews having a continuous breeding range and a pattern of parallel migration, all three approaches identified groups with different population trends, movement behaviours and habitat selection across the annual cycle, suggesting these are meaningful ecological groups. The expert and clustering approaches resulted in similar group structure, strong estimates of migratory connectivity (measured as MC = 0.64 across seasons), movement behaviour and habitat selection; however, the expert approach identified an additional divide between the easternmost grouping, which revealed strongly negative population trends in the group occupying the Chihuahuan desert during the stationary nonbreeding season. In contrast, the flyway delineation resulted in weaker estimates of migratory connectivity, marginal differences in population trends and less between-group differences in movement behaviour and habitat selection. 4. Synthesis and applications . Using measurements of migratory connectivity in concert with expert opinion can define ecologically distinct groups for wildlife management that differ in the environmental conditions they experience across seasons of the annual cycle, which is a key component for understanding and reversing declines of migratory species.

Journal of Applied Ecology

Filling knowledge gaps in a threatened shorebird flyway through satellite tracking

Satellite‐based technologies that track individual animal movements enable the mapping of their spatial and temporal patterns of occurrence. This is particularly useful in poorly studied or remote regions where there is a need for the rapid gathering of relevant ecological knowledge to inform management actions. One such region is East Asia, where many intertidal habitats are being degraded at unprecedented rates and shorebird populations relying on these habitats show rapid declines. We examine the utility of satellite tracking to accelerate the identification of coastal sites of conservation importance in the East Asian–Australasian Flyway. In 2015–2017, we used solar‐powered satellite transmitters to track the migration of 32 great knots ( Calidris tenuirostris ), an “Endangered” shorebird species widely distributed in the Flyway and fully dependent on intertidal habitats for foraging during the non‐breeding season. From the great knot tracks, a total of 92 stopping sites along the Flyway were identified. Surprisingly, 63% of these sites were not known as important shorebird sites before our study; in fact, every one of the tracked individuals used sites that were previously unrecognized. Site knowledge from on‐ground studies in the Flyway is most complete for the Yellow Sea and generally lacking for Southeast Asia, Southern China and Eastern Russia. Synthesis and applications . Satellite tracking highlighted coastal habitats that are potentially important for shorebirds but lack ecological information and conservation recognition, such as those in Southern China and Southeast Asia. At the same time, the distributional data of tracked individuals can direct on‐ground surveys at the lesser known sites to collect information on bird numbers and habitat characteristics. To recognize and subsequently protect valuable coastal habitats, filling knowledge gaps by integrating bird tracking with ground‐based methods should be prioritized.

East Asian-Australasian Flyway

FORUM: Effective management of ecological resilience – are we there yet?

Ecological resilience is developing into a credible paradigm for policy development and environmental management for preserving natural capital in a rapidly changing world. However, resilience emerges from complex interactions, limiting the translation of theory into practice. Main limitations include the following: (i) difficulty in quantification and detection of changes in ecological resilience, (ii) a lack of empirical evidence to support preventative or proactive management and (iii) difficulties in managing processes operating across socio-ecological systems that vary in space and time. We highlight recent research with the potential to address these limitations including new and/or improved indicators of resilience and tools to assess scale as a driver of resilience. Synthesis and applications . Effective resilience-based management must be adaptive in nature. To support this, we propose an operational model using resilience-based iterative management actions operating across scales.

Journal of Applied Ecology

The transition from resistance to acceptance: Managing a marine invasive species in a changing world

Marine invasive species can transform coastal ecosystems, yet mitigating their effects can be difficult, and even impractical. Often, marine invasive species are managed at poorly matched spatial scales, and at the same time, rates of spread and establishment are increasing under climate change and can outpace resources available for population suppression. These circumstances challenge traditional conservation goals of maintaining a historic environmental state, especially for a species like the European green crab ( Carcinus maenas ), a formidable invader with few examples of successful long-term removal programs. A management paradigm where decision alternatives include resisting or accepting a new ecological trajectory may be needed. We apply mathematical concepts from decision theory to develop a quantitative framework for navigating management decisions in this new resist-accept paradigm. We develop a model of European green crab growth, removal and colonization, and we find optimal levels of removal effort that minimize both ecological change and removal cost. We establish a benchmark of colonization pressure at which green crab density becomes decoupled from a decision maker's actions, such that population control can no longer shape the invasion trajectory. For informing the decision boundary between resistance and acceptance, our results highlight that a decision maker's understanding of how removal cost scales with removal effort is more important than understanding the density-impact relationship. We show that assuming stationary system dynamics can result in sub-optimal levels of species removal effort, highlighting the importance of developing anticipatory management strategies by accounting for non-stationary dynamics. Policy implications . For marine invasive species that can disperse across long distances and recolonize rapidly after removal, the focus of conservation policy should shift away from understanding how to resist change to understanding when to stop resisting change. Navigating this decision problem involves trade-offs among competing objectives, highlighting the need for structured approaches to elicit objective weights that reflect the values of the decision maker. For natural resource managers facing possible ecosystem transformation, this decision framework can enable proactive and strategic decisions made under uncertainty in a changing world.

Journal of Applied Ecology

Reanalysis ignores pertinent data, includes inappropriate observations, and disregards realities of applied ecology: Response to Huso and Dalthorp (2023)

1) We recently demonstrated efficacy of automated curtailment of wind turbines in reducing fatalities of eagles at a study site in Wyoming, USA. Huso and Dalthorp criticize our work, asserting that there are several ‘major errors’ that render our previous work as providing ‘no meaningful support’ for automated curtailment. As we show here, our data do indeed provide support for the efficacy of automated curtailment. 2) The purported major errors they identify include: having only 1 year of post-treatment data, having only one control and one treatment site, using the control site for inference, choosing an arbitrary demarcation date between the ‘before’ and ‘after’ periods, analysing the number, rather than the rate, of fatalities, and not including data they consider pertinent. These claims are unsupported because all result from misreading our study, ignoring realities of causal inference in field ecology, or are matters of preference. 3) Most importantly, we did not fail to include pertinent data and we provided sufficient criteria in the manuscript to explain inclusion of carcasses in our study. As stated in our previous study, the eagle carcasses we did not include were found either outside of search areas or incidentally, and thus did not meet criteria for inclusion. Further, Huso and Dalthorp present a standard for spatial and temporal replication that is inconsistent with their own recent work and many other studies in this field. Finally, their reanalysis of our data ignores the control site while including inappropriate data and thus is not suitable for inference. 4) We appreciate the unusually thorough critique of our work provided by Huso and Dalthorp. Despite the major errors in their argument, this discussion provides a platform for further evaluation of our original work. 5) Synthesis and applications . We show that incorporating control site data and using our a priori demarcation of before and after periods into Huso and Dalthorp's reanalysis returns meaningful support for automated curtailment. We also direct the reader to an updated analysis that uses components of the approach Huso and Dalthorp suggest, along with additional data and a number of other analytical improvements, to validate and strengthen our original conclusions.

Journal of Applied Ecology

Evaluation of trap capture in a geographically closed population of brown treesnakes on Guam

1. Open population mark-recapture analysis of unbounded populations accommodates some types of closure violations (e.g. emigration, immigration). In contrast, closed population analysis of such populations readily allows estimation of capture heterogeneity and behavioural response, but requires crucial assumptions about closure (e.g. no permanent emigration) that are suspect and rarely tested empirically. 2. In 2003, we erected a double-sided barrier to prevent movement of snakes in or out of a 5-ha semi-forested study site in northern Guam. This geographically closed population of >100 snakes was monitored using a series of transects for visual searches and a 13 ?? 13 trapping array, with the aim of marking all snakes within the site. Forty-five marked snakes were also supplemented into the resident population to quantify the efficacy of our sampling methods. We used the program mark to analyse trap captures (101 occasions), referenced to census data from visual surveys, and quantified heterogeneity, behavioural response, and size bias in trappability. Analytical inclusion of untrapped individuals greatly improved precision in the estimation of some covariate effects. 3. A novel discovery was that trap captures for individual snakes consisted of asynchronous bouts of high capture probability lasting about 7 days (ephemeral behavioural effect). There was modest behavioural response (trap happiness) and significant latent (unexplained) heterogeneity, with small influences on capture success of date, gender, residency status (translocated or not), and body condition. 4. Trapping was shown to be an effective tool for eradicating large brown treesnakes Boiga irregularis (>900 mm snout-vent length, SVL). 5. Synthesis and applications. Mark-recapture modelling is commonly used by ecological managers to estimate populations. However, existing models involve making assumptions about either closure violations or response to capture. Physical closure of our population on a landscape scale allowed us to determine the relative importance of covariates influencing capture probability (body size, trappability periods, and latent heterogeneity). This information was used to develop models in which different segments of the population could be assigned different probabilities of capture, and suggests that modelling of open populations should incorporate easily measured, but potentially overlooked, parameters such as body size or condition. ?? 2008 The Authors.

Journal of Applied Ecology

The application of oyster reefs in shoreline protection: Are we over‐engineering for an ecosystem engineer?

Oyster reef living shorelines have been proposed as an effective alternative to traditional coastal defence structures (e.g. bulkheads, breakwaters), with the benefit that they may keep pace with sea‐level rise and provide co‐benefits, such as habitat provision. However, there remains uncertainty about the effectiveness of shoreline protection provided by oyster reefs, which limits their broader application. We draw evidence from studies along the east and gulf coasts of the United States, where much research and implementation of oyster reef restoration has occurred, to better define the existing gaps in our understanding of the use of restored oyster reefs for shoreline protection. We find potential disconnects between ecological and engineering functions of reefs. In response, we outline how engineering and ecological principles are used in the design of oyster reef living shorelines and highlight knowledge gaps where an integration of these disciplines will lead to their more effective application. Synthesis and applications . This work highlights the necessary steps to advance the application of oyster reef living shorelines. Importantly, future research should focus on appropriate designs and conditions needed for these structures to effectively protect our coasts from erosion, while supporting a sustainable oyster population, thereby providing actionable nature‐based alternatives for coastal defence to diverse end‐users.

East Coast of Continental United States

Improving estimation of flight altitude in wildlife telemetry studies

Altitude measurements from wildlife tracking devices, combined with elevation data, are commonly used to estimate the flight altitude of volant animals. However, these data often include measurement error. Understanding this error may improve estimation of flight altitude and benefit applied ecology. There are a number of different approaches that have been used to address this measurement error. These include filtering based on GPS data, filtering based on behaviour of the study species, and use of state-space models to correct measurement error. The effectiveness of these approaches is highly variable. Recent studies have based inference of flight altitude on misunderstandings about avian natural history and technical or analytical tools. In this Commentary, we discuss these misunderstandings and suggest alternative strategies both to resolve some of these issues and to improve estimation of flight altitude. These strategies also can be applied to other measures derived from telemetry data. Synthesis and applications. Our Commentary is intended to clarify and improve upon some of the assumptions made when estimating flight altitude and, more broadly, when using GPS telemetry data. We also suggest best practices for identifying flight behaviour, addressing GPS error, and using flight altitudes to estimate collision risk with anthropogenic structures. Addressing the issues we describe would help improve estimates of flight altitude and advance understanding of the treatment of error in wildlife telemetry studies.

Journal of Applied Ecology

Quantifying resilience

The biosphere is under unprecedented pressure, reflected in rapid changes in our global ecological, social, technological and economic systems. In many cases, ecological and social systems can adapt to these changes over time, but when a critical threshold is surpassed, a system under stress can undergo catastrophic change and reorganize into a different state. The concept of resilience, introduced more than 40 years ago in the ecological sciences, captures the behaviour of systems that can occur in alternative states. The original definition of resilience forwarded by Holling ( 1973 ) is still the most useful. It defines resilience as the amount of disturbance that a system can withstand before it shifts into an alternative stable state. The idea of alternative stable states has clear and profound implications for ecological management. Coral reefs, for example, are high-diversity systems that provide key ecosystem services such as fisheries and coastal protection. Human impacts are causing significant, ongoing reef degradation, and many reefs have shifted from coral- to algal-dominated states in response to anthropogenic pressures such as elevated water temperatures and overfishing. Understanding and differentiating between the factors that help maintain reefs in coral-dominated states vs. those that facilitate a shift to an undesired algal-dominated state is a critical step towards sound management and conservation of these, and other, important social&ndash;ecological systems. Resilience has gained popularity among both academicians and laypeople, as a term meant to describe a systems&rsquo; ability to withstand disturbance. Resilience has become a buzzword in the last decade, as shown by its increasing appearance in calls for research proposals and scientific citation data bases. The term resilience has in many cases lost the clarity of the original definition and in fact is frequently used in a manner in direct opposition to the original definition. Many current uses of the concept are loose and incorrect. The term is becoming increasingly used in a normative sense (Brand & Jax 2007 ), as if resilience were a desirable quality of systems. However, even systems in highly undesirable states, such as macro-algae dominated reefs, or city cores in poverty traps, may be highly resilient, which is to say they withstand attempts to transform them into different (desirable) states. Operationalizing the concept of resilience for application and management has been difficult. Misuse of the term can have significant negative impacts, because resilience is being used to help guide responses to natural disasters and to assess the sustainability of ecosystems and urban systems and has been driving international research priorities. Resilience has been argued to be a basic emergent property of systems, a process or a rate. We focus on the original concept as described by Holling, which is that of an emergent system property; when a system is in a desirable state and managers wish to enhance resilience, or when the system is in an undesirable state and managers wish to erode resilience and foster a transformation to an alternative state. Fostering or eroding resilience is a process. When a system is perturbed but resilience is not exceeded, then the recovery can be measured as a rate. Several frameworks to operationalize resilience have been proposed. A decade ago, a special feature focused on quantifying resilience was published in the journal Ecosystems (Carpenter, Westley & Turner 2005 ). The approach there was towards identifying surrogates of resilience, but few of the papers proposed quantifiable metrics. Consequently, many ecological resilience frameworks remain vague and difficult to quantify, a problem that this special feature aims to address. However, considerable progress has been made during the last decade (e.g. Pope, Allen & Angeler 2014 ). Although some argue that resilience is best kept as an unquantifiable, vague concept (Quinlan et al . 2016 ), to be useful for managers, there must be concrete guidance regarding how and what to manage and how to measure success (Garmestani, Allen & Benson 2013 ; Spears et al . 2015 ). Ideas such as &lsquo;resilience thinking&rsquo; have utility in helping stakeholders conceptualize their systems, but provide little guidance on how to make resilience useful for ecosystem management, other than suggesting an ambiguous, Goldilocks approach of being just right (e.g. diverse, but not too diverse; connected, but not too connected). Here, we clarify some prominent resilience terms and concepts, introduce and synthesize the papers in this special feature on quantifying resilience and identify core unanswered questions related to resilience.

Journal of Applied Ecology

Occupancy estimation and the closure assumption

1. Recent advances in occupancy estimation that adjust for imperfect detection have provided substantial improvements over traditional approaches and are receiving considerable use in applied ecology. To estimate and adjust for detectability, occupancy modelling requires multiple surveys at a site and requires the assumption of 'closure' between surveys, i.e. no changes in occupancy between surveys. Violations of this assumption could bias parameter estimates; however, little work has assessed model sensitivity to violations of this assumption or how commonly such violations occur in nature. 2. We apply a modelling procedure that can test for closure to two avian point-count data sets in Montana and New Hampshire, USA, that exemplify time-scales at which closure is often assumed. These data sets illustrate different sampling designs that allow testing for closure but are currently rarely employed in field investigations. Using a simulation study, we then evaluate the sensitivity of parameter estimates to changes in site occupancy and evaluate a power analysis developed for sampling designs that is aimed at limiting the likelihood of closure. 3. Application of our approach to point-count data indicates that habitats may frequently be open to changes in site occupancy at time-scales typical of many occupancy investigations, with 71% and 100% of species investigated in Montana and New Hampshire respectively, showing violation of closure across time periods of 3 weeks and 8 days respectively. 4. Simulations suggest that models assuming closure are sensitive to changes in occupancy. Power analyses further suggest that the modelling procedure we apply can effectively test for closure. 5. Synthesis and applications. Our demonstration that sites may be open to changes in site occupancy over time-scales typical of many occupancy investigations, combined with the sensitivity of models to violations of the closure assumption, highlights the importance of properly addressing the closure assumption in both sampling designs and analysis. Furthermore, inappropriately applying closed models could have negative consequences when monitoring rare or declining species for conservation and management decisions, because violations of closure typically lead to overestimates of the probability of occurrence.

Montana;New Hampshire

Habitat drives dispersal and survival of translocated juvenile desert tortoises

1.In spite of growing reliance on translocations in wildlife conservation, translocation efficacy remains inconsistent. One factor that can contribute to failed translocations is releasing animals into poor quality or otherwise inadequate habitat. 2.Here we used a targeted approach to test the relationship of habitat features to post-translocation dispersal and survival of juvenile Mojave desert tortoises Gopherus agassizii . 3.We selected three habitat characteristics&mdash;rodent burrows, substrate texture (prevalence and size of rocks), and washes (ephemeral river beds)&ndash;that are tied to desert tortoise ecology. At the point of release, we documented rodent burrow abundance, substrate texture, and wash presence and analysed their relationship to maximum dispersal. We also documented relative use by each individual for each habitat characteristic and analysed their relationships with survival and fatal encounters with a predator in the first year after release. 4.In general, the presence of refugia or other areas that enabled animals to avoid detection, such as burrows and substrate, decreased overall mortality as well as predator-mediated mortality. The presence of washes and substrate that enhanced the tortoises&rsquo; ability to avoid detection also associated with reduced dispersal away from the release site. These results indicate an important role for all three measured habitat characteristics in driving dispersal, survival, or fatal encounters with a predator in the first year after translocation. 5. Synthesis and applications . Resource managers using translocations as a conservation tool should prioritize acquiring data linking habitat to fitness. In particular, for species that depend on avoiding detection, refuges such as burrows and habitat that improved concealment had notable ability to improve survival and dispersal. Our study on juvenile Mojave desert tortoises showed that refuge availability or the distributions of habitat appropriate for concealment are important considerations for identifying translocation sites for species highly dependent on crypsis, camouflage, or other forms of habitat matching.

Journal of Applied Ecology