Using the R package popharvest to assess the sustainability of offtake in birds.

The R package popharvest was designed to help assess the sustainability of offtake in birds when only limited demographic information is available. In this article, we describe some basics of harvest theory and then discuss several considerations when using the different approaches in popharvest to assess whether observed harvests are unsustainable. Throughout, we emphasize the importance of distinguishing between the scientific and policy aspects of managing offtake. The principal product of popharvest is a sustainable harvest index (SHI), which can indicate whether the harvest is unsustainable but not the converse. SHI is estimated based on a simple, scalar model of logistic population growth, whose parameters may be estimated using limited knowledge of demography. Uncertainty in demography leads to a distribution of SHI values and it is the purview of the decision-maker to determine what amounts to an acceptable risk when failing to reject the null hypothesis of sustainability. The attitude toward risk, in turn, will likely depend on the decision-maker's objective(s) in managing offtake. The management objective as specified in popharvest is a social construct, informed by biology, but ultimately it is an expression of social values that usually vary among stakeholders. We therefore suggest that any standardization of criteria for management objectives in popharvest will necessarily be subjective and, thus, hard to defend in diverse decision-making situations. Because of its ease of use, diverse functionalities, and a minimal requirement of demographic information, we expect the use of popharvest to become widespread. Nonetheless, we suggest that while popharvest provides a useful platform for rapid assessments of sustainability, it cannot substitute for sufficient expertise and experience in harvest theory and management.

Age-specific survival rates, causes of death, and allowable take of golden eagles in the western United States.

In the United States, the Bald and Golden Eagle Protection Act prohibits take of golden eagles (Aquila chrysaetos) unless authorized by permit, and stipulates that all permitted take must be sustainable. Golden eagles are unintentionally killed in conjunction with many lawful activities (e.g., electrocution on power poles, collision with wind turbines). Managers who issue permits for incidental take of golden eagles must determine allowable take levels and manage permitted take accordingly. To aid managers in making these decisions in the western United States, we used an integrated population model to obtain estimates of golden eagle vital rates and population size, and then used those estimates in a prescribed take level (PTL) model to estimate the allowable take level. Estimated mean annual survival rates for golden eagles ranged from 0.70 (95% credible interval = 0.66-0.74) for first-year birds to 0.90 (0.88-0.91) for adults. Models suggested a high proportion of adult female golden eagles attempted to breed and breeding pairs fledged a mean of 0.53 (0.39-0.72) young annually. Population size in the coterminous western United States has averaged ~31,800 individuals for several decades, with λ = 1.0 (0.96-1.05). The PTL model estimated a median allowable take limit of ~2227 (708-4182) individuals annually given a management objective of maintaining a stable population. We estimate that take averaged 2572 out of 4373 (59%) deaths annually, based on a representative sample of transmitter-tagged golden eagles. For the subset of golden eagles that were recovered and a cause of death determined, anthropogenic mortality accounted for an average of 74% of deaths after their first year; leading forms of take over all age classes were shooting (~670 per year), collisions (~611), electrocutions (~506), and poisoning (~427). Although observed take overlapped the credible interval of our allowable take estimate and the population overall has been stable, our findings indicate that additional take, unless mitigated for, may not be sustainable. Our analysis demonstrates the utility of the joint application of integrated population and prescribed take level models to management of incidental take of a protected species.

Evaluation of a two-season banding program to estimate and model migratory bird survival.

The management of North American waterfowl is predicated on long-term, continental-scale banding implemented prior to the hunting season (i.e., July-September) and subsequent reporting of bands recovered by hunters. However, single-season banding and encounter operations have a number of characteristics that limit their application to estimating demographic rates and evaluating hypothesized limiting factors throughout the annual cycle. We designed and implemented a two-season banding program for American Black Ducks (Anas rubripes), Mallards (A. platyrhynchos), and hybrids in eastern North America to evaluate potential application to annual life cycle conservation and sport harvest management. We assessed model fit and compared estimates of annual survival among data types (i.e., pre-hunting season only [July-September], post-hunting season only [January-March], and two-season [pre- and post-hunting season]) to evaluate model assumptions and potential application to population modeling and management. There was generally high agreement between estimates of annual survival derived using two-season and pre-season only data for all age and sex cohorts. Estimates of annual survival derived from post-season banding data only were consistently higher for adult females and juveniles of both sexes. We found patterns of seasonal survival varied by species, age, and to a lesser extent, sex. Hunter recovered birds exhibited similar spatial distributions regardless of banding season suggesting banded samples were from the same population. In contrast, goodness-of-fit tests suggest this assumption was statistically violated in some regions and years. We conclude that estimates of seasonal and annual survival for Black Ducks and Mallards based on the two-season banding program are valid and accurate based on model fit statistics, similarity in survival estimates across data and models, and similarities in the distribution of recoveries. The two-season program provides greater precision and insight into the survival process and will improve the ability of researchers and managers to test competing hypotheses regarding population regulation resulting in more effective management.

Detection probabilities of bird carcasses along sandy beaches and marsh edges in the northern Gulf of Mexico.

We estimated detection probabilities of bird carcasses along sandy beaches and in marsh edge habitats in the northern Gulf of Mexico to help inform models of bird mortality associated with the Deepwater Horizon oil spill. We also explored factors that may influence detection probability, such as carcass size, amount of scavenging, location on the beach, habitat type, and distance into the marsh. Detection probability for medium-sized carcasses (200-500 g) ranged from 0.82 (SE = 0.09) to 0.93 (SE = 0.04) along sandy beaches. Within sandy beaches, we found that intact/slightly scavenged carcasses were easier to detect than heavily scavenged ones and did not find strong effects of location on the beach on detection probability. We estimated detection rate for each combination of scavenging state, carcass size, and position along sandy beaches. In marsh edge habitats, detection ranged from 0.04 (SE = 0.04) to 0.86 (SE = 0.10), with detection rates rapidly increasing from small (< 200 g) to medium carcass sizes and leveling off between medium and extra-large (> 1000 g) carcasses regardless of vegetation type (Spartina or Phragmites). Carcasses of all sizes were generally harder to locate in Spartina-dominated marshes than in Phragmites-dominated ones. A subset of the data for which we could adequately assess the effect of distance into the marsh indicated that detection rates generally declined the farther a carcass was into marsh vegetation. Based on power analyses, our ability to identify predictors that influence detection rates would be higher with larger numbers of carcasses, greater numbers of search trials per carcass, or more balanced sampling distributions across predictor values.

Persistence of avian carcasses on sandy beaches and marsh edges in the northern Gulf of Mexico.

As part of the natural resource damage assessment for the Deepwater Horizon oil spill, a mathematical model was used to estimate the total number of bird carcasses deposited on shorelines based on the number of carcasses collected and adjustment factors such as detection probability and carcass persistence. Studies of carcass persistence occurred along sandy beaches and marsh edges in the northern Gulf of Mexico to obtain site-specific inputs for the model. We estimated persistence rates for these habitat types and evaluated the influence on persistence of carcass size, location of the carcass on the beach, dominant vegetation type in the marsh, carcass distance into marsh vegetation, and length of time a carcass was stranded on a shoreline. The length of time stranded had the greatest influence on persistence in both habitat types, with persistence initially relatively low and increasing logarithmically. Carcass size and position were weakly influential on sandy beaches. Carcass size had stronger influences along marsh edges, and marsh habitat type also affected persistence. We found evidence of a positive relationship between distance into the marsh and persistence during the first 24 h after carcass deployment.

Evaluation of harvest and information needs for North American sea ducks.

Wildlife managers routinely seek to establish sustainable limits of sport harvest or other regulated forms of take while confronted with considerable uncertainty. A growing body of ecological research focuses on methods to describe and account for uncertainty in management decision-making and to prioritize research and monitoring investments to reduce the most influential uncertainties. We used simulation methods incorporating measures of demographic uncertainty to evaluate risk of overharvest and prioritize information needs for North American sea ducks (Tribe Mergini). Sea ducks are popular game birds in North America, yet they are poorly monitored and their population dynamics are poorly understood relative to other North American waterfowl. There have been few attempts to assess the sustainability of harvest of North American sea ducks, and no formal harvest strategy exists in the U.S. or Canada to guide management. The popularity of sea duck hunting, extended hunting opportunity for some populations (i.e., special seasons and/or bag limits), and population declines have led to concern about potential overharvest. We used Monte Carlo simulation to contrast estimates of allowable harvest and observed harvest and assess risk of overharvest for 7 populations of North American sea ducks: the American subspecies of common eider (Somateria mollissima dresseri), eastern and western populations of black scoter (Melanitta americana) and surf scoter (M. perspicillata), and continental populations of white-winged scoter (M. fusca) and long-tailed duck (Clangula hyemalis). We combined information from empirical studies and the opinions of experts through formal elicitation to create probability distributions reflecting uncertainty in the individual demographic parameters used in this assessment. Estimates of maximum growth (rmax), and therefore of allowable harvest, were highly uncertain for all populations. Long-tailed duck and American common eider appeared to be at high risk of overharvest (i.e., observed harvest < allowable harvest in 5-7% and 19-26% of simulations, respectively depending on the functional form of density dependence), whereas the other populations appeared to be at moderate risk to low risk (observed harvest < allowable harvest in 22-68% of simulations, again conditional on the form of density dependence). We also evaluated the sensitivity of the difference between allowable and observed harvest estimates to uncertainty in individual demographic parameters to prioritize information needs. We found that uncertainty in overall fecundity had more influence on comparisons of allowable and observed harvest than adult survival or observed harvest for all species except long-tailed duck. Although adult survival was characterized by less uncertainty than individual components of fecundity, it was identified as a high priority information need given the sensitivity of growth rate and allowable harvest to this parameter. Uncertainty about population size was influential in the comparison of observed and allowable harvest for 5 of the 6 populations where it factored into the assessment. While this assessment highlights a high degree of uncertainty in allowable harvest, it provides a framework for integration of improved data from future research and monitoring. It could also serve as the basis for harvest strategy development as management objectives and regulatory alternatives are specified by the management community.

State-Dependent Resource Harvesting with Lagged Information about System States.

Markov decision processes (MDPs), which involve a temporal sequence of actions conditioned on the state of the managed system, are increasingly being applied in natural resource management. This study focuses on the modification of a traditional MDP to account for those cases in which an action must be chosen after a significant time lag in observing system state, but just prior to a new observation. In order to calculate an optimal decision policy under these conditions, possible actions must be conditioned on the previous observed system state and action taken. We show how to solve these problems when the state transition structure is known and when it is uncertain. Our focus is on the latter case, and we show how actions must be conditioned not only on the previous system state and action, but on the probabilities associated with alternative models of system dynamics. To demonstrate this framework, we calculated and simulated optimal, adaptive policies for MDPs with lagged states for the problem of deciding annual harvest regulations for mallards (Anas platyrhynchos) in the United States. In this particular example, changes in harvest policy induced by the use of lagged information about system state were sufficient to maintain expected management performance (e.g. population size, harvest) even in the face of an uncertain system state at the time of a decision.

A flexible Bayesian hierarchical approach for analyzing spatial and temporal variation in the fecal corticosterone levels in birds when there is imperfect knowledge of individual identity.

Population cycles have long interested biologists. The ruffed grouse, Bonasa umbellus, is one such species whose populations cycle over most of their range. Thus, much effort has been expended to understand the mechanisms that might control cycles in this and other species. Corticosterone metabolites are widely used in studies of animals to measure physiological stress. We evaluated corticosterone metabolites in feces of territorial male grouse as a potential tool to study mechanisms governing grouse cycles. However, like most studies of corticosterone in wild animals, we did not know the identity of all individuals for which we had fecal samples. This presented an analytical problem that resulted in either pseudoreplication or confounding. Therefore, we derived an analytical approach that accommodated for uncertainty in individual identification. Because we had relatively low success capturing birds, we estimated turnover probabilities of birds on territorial display sites based on capture histories of a limited number of birds we captured. Hence, we developed a study design and modeling approach to quantify variation in corticosterone levels among individuals and through time that would be applicable to any field study of corticosterone in wild animals. Specifically, we wanted a sampling design and model that was flexible enough to partition variation among individuals, spatial units, and years, while incorporating environmental covariates that would represent potential mechanisms. We conducted our study during the decline phase of the grouse cycle and found high variation among corticosterone samples (11.33-443.92 ng/g [x=113.99 ng/g, SD=69.08, median=99.03 ng/g]). However, there were relatively small differences in corticosterone levels among years, but levels declined throughout each breeding season, which was opposite our predictions for stress hormones correlating with a declining population. We partitioned the residual variation into site, bird, and repetition (i.e., multiple samples collected from the same bird on the same day). After accounting for years and three general periods within breeding seasons, 42% of the residual variation among observations was attributable to differences among individual birds. Thus, we attribute little influence of site on stress level of birds in our study, but disentangling individual from site effects is difficult because site and bird are confounded. Our model structures provided analytical approaches for studying species having different ecologies. Our approach also demonstrates that even incomplete information on individual identity of birds within samples is useful for analyzing these types of data.