Management and Recovery Options for Ural River Beluga Sturgeon

Abstract: Management of declining fisheries of anadromous species sometimes relies heavily on supplementation of populations with captive breeding, despite evidence that captive breeding can have negative consequences and may not address the root cause of decline. The beluga sturgeon (Huso huso), a species threatened by the market for black caviar and reductions in habitat quality, is managed through harvest control and hatchery supplementation, with an emphasis on the latter. We used yield per recruit and elasticity analyses to evaluate the population status and current levels of fishing and to identify the life‐history stages that are the best targets for conservation of beluga of the Ural River. Harvest rates in recent years were four to five times higher than rates that would sustain population abundance. Sustainable rates of fishing mortality are similar to those for other long‐lived marine species such as sharks and mammals. Yield per recruit, which is maximized if fish are first harvested at age 31 years, would be greatly enhanced by raising minimum size limits or reducing illegal take of subadults. Improving the survival of subadult and adult females would increase population productivity by 10 times that achieved by improving fecundity and survival from egg to age 1 year (i.e., hatchery supplementation). These results suggest that reducing mortality of subadults and adult wild fish is a more effective conservation strategy than hatchery supplementation. Because genetics is not factored into hatchery management practices, supplementation may even reduce the viability of the beluga sturgeon.     

Life History as a Predictor of Salamander Recovery Rate from Timber Harvest in Southern Appalachian Forests,U.S.A

Forest management often represents a balance between social, economic, and ecological objectives. In the eastern United States, numerous studies have established that terrestrial salamander populations initially decline in abundance following timber harvest, yet the large‐scale and long‐term consequences are relatively unknown. We used count data from terrestrial survey points to examine the relation between salamander abundance and historic timber harvest while accounting for imperfect detection of individuals. Overall, stream‐ and terrestrial‐breeding salamanders appeared to differ by magnitude of population decline, rate of population recovery, and extent of recolonization from surrounding forest. Specifically, estimated abundance of both species groups was positively associated with stand age and recovery rates were predicted to increase over time for red‐legged salamanders (Plethodon shermani) and decrease in stream‐breeding species. Abundance of stream‐breeding salamanders was predicted to reach a peak by 100 years after timber harvest, and the population growth rate of red‐legged salamanders was predicted to undergo a significant increase 100 years after harvest. Estimated abundance of stream‐breeding salamanders in young forest stands was also negatively associated with the distance to adjacent forest, a result that suggests immigration has a role in the recovery of these species. Our results indicate that salamander abundance in young forest stands may be only modestly lower than in more mature forest but that full recovery from timber harvest may take a substantial amount of time and that species life history may affect patterns of recovery. Historia de Vida como un Vaticinador de la Tasa de Recuperación de una Salamandra a la Colecta de Madera en los Bosques del Sur de los Apalaches, E.U.A     

Conserving spawning stocks through harvest slot limits and no-take protected areas

The key to the conservation of harvested species is the maintenance of reproductive success. Yet for many marine species large, old, individuals are targeted despite their disproportionate contribution to reproduction. We hypothesized that a combination of no-take marine protected areas (MPAs) and harvest slot limits (maximum and minimum size limits) would result in the conservation of large spawning individuals under heavy harvest. We tested this approach under different harvest intensities with a 2-sex, stage-structured metapopulation model for the Caribbean spiny lobster (Panulirus argus). P. argus is intensively harvested in the Caribbean, and in many localities large, mature individuals no longer exist. No-take MPAs and harvest slot limits combined, rebuilt and maintained large mature individuals even under high harvest pressure. The most conservative model (a 30% MPA and harvest slot limit of 75–105 mm) increased spawner abundance by 5.53E¹² compared with the fishing status quo at the end of 30 years. Spawning stock abundance also increased by 2.76–9.56E¹² individuals at a high harvest intensity over 30 years with MPAs alone. Our results demonstrate the potential of MPAs and harvest slot limits for the conservation of large breeding individuals in some marine and freshwater environments. Decisions on which management strategy best suits a fishery, however, requires balancing what is ecologically desirable with what is economically and socially feasible.     

Sex ratio and optimal harvest of canvasback ducks,A model

The North American population of canvasback ducks (Aythya valisineria) exhibits extreme distortion of the sex ratio in favor of males. This paper describes a model which accounts for this pattern by relatively heavier female mortality in both the breeding and nonbreeding seasons. The density-dependence of winter mortality leads to the conclusion that the observed sex ratio depresses total population numbers. Variation in nesting success is shown to influence sex ratios and strongly depress population numbers. Because a standard harvest scheme can be demonstrated to severely depress the numbers of ducks, an alternative graduated or weighted harvest procedure is recommended.     

Defining trade-offs among conservation, profitability, and food security in the California current bottom-trawl fishery

Although it is recognized that marine wild-capture fisheries are an important source of food for much of the world, the cost of sustainable capture fisheries to species diversity is uncertain, and it is often questioned whether industrial fisheries can be managed sustainably. We evaluated the trade-off among sustainable food production, profitability, and conservation objectives in the groundfish bottom-trawl fishery off the U.S. West Coast, where depletion (i.e., reduction in abundance) of six rockfish species (Sebastes) is of particular concern. Trade-offs are inherent in this multispecies fishery because there is limited capacity to target species individually. From population models and catch of 34 stocks of bottom fish, we calculated the relation between harvest rate, long-term yield (i.e., total weight of fish caught), profit, and depletion of each species. In our models, annual ecosystem-wide yield from all 34 stocks was maximized with an overall 5.4% harvest rate, but profit was maximized at a 2.8% harvest rate. When we reduced harvest rates to the level (2.2% harvest rate) at which no stocks collapsed (<10% of unfished levels), biomass harvested was 76% of the maximum sustainable yield and profit 89% of maximum. A harvest rate under which no stocks fell below the biomass that produced maximum sustainable yield (1% harvest rate), resulted in 45% of potential yield and 67% of potential profit. Major reductions in catch in the late 1990s led to increase in the biomass of the most depleted stocks, but this rebuilding resulted in the loss of >30% of total sustainable yield, whereas yield lost from stock depletion was 3% of total sustainable yield. There are clear conservation benefits to lower harvest rates, but avoiding overfishing of all stocks in a multispecies fishery carries a substantial cost in terms of lost yield and profit.     

Maximizing Offspring Production While Maintaining Genetic Diversity in Supplemental Breeding Programs of Highly Fecund Managed Species

Abstract:  Supplemental breeding is an intensive population management strategy wherein adults are captured from nature and spawned in controlled settings, and the resulting offspring are later released into the wild. To be effective, supplemental breeding programs require crossing strategies that maximize offspring production while maintaining genetic diversity within each supplemental year class. We used computer simulations to assess the efficacy of different mating designs to jointly maximize offspring production and maintain high levels of genetic diversity (as measured by the effective population size) under a variety of biological conditions particularly relevant to species with high fecundity and external fertilization, such as many fishes. We investigated four basic supplemental breeding designs involving either monogamous pairings or complete factorial designs (in which every female is mated to every male and vice versa), each with or without the added stipulation that all breeders contribute equally to the total reproductive output. In general, complete factorial designs that did not equalize parental contributions came closest to the goal of maximizing offspring production while still maintaining relatively large effective population sizes. Next, we estimated the effective population size of 10 different supplemental year classes within the breeding program of the robust redhorse (Moxostoma robustum). Two year classes failed to produce progeny, whereas successful year classes used partial factorial designs to realize effective sizes ranging from 2 to 26 individuals. On average, a complete factorial design could increase the effective size of each robust redhorse supplemental year class by 19%.     

Modeling the Effects of Trophy Selection and Environmental Disturbance on a Simulated Population of African Lions

Abstract:  Tanzania is a premier destination for trophy hunting of African lions (Panthera leo) and is home to the most extensive long-term study of unhunted lions. Thus, it provides a unique opportunity to apply data from a long-term field study to a conservation dilemma: How can a trophy-hunted species whose reproductive success is closely tied to social stability be harvested sustainably? We used an individually based, spatially explicit, stochastic model, parameterized with nearly 40 years of behavioral and demographic data on lions in the Serengeti, to examine the separate effects of trophy selection and environmental disturbance on the viability of a simulated lion population in response to annual harvesting. Female population size was sensitive to the harvesting of young males (≥3 years), whereas hunting represented a relatively trivial threat to population viability when the harvest was restricted to mature males (≥6 years). Overall model performance was robust to environmental disturbance and to errors in age assessment based on nose coloration as an index used to age potential trophies. Introducing an environmental disturbance did not eliminate the capacity to maintain a viable breeding population when harvesting only older males, and initially depleted populations recovered within 15–25 years after the disturbance to levels comparable to hunted populations that did not experience a catastrophic event. These results are consistent with empirical observations of lion resilience to environmental stochasticity .     

A population viability analysis for the Island Fox on Santa Catalina Island,California

The island fox (Urocyon littoralis) on Santa Catalina Island is among the most imperiled species on the Channel Islands due to a recent outbreak of canine distemper virus (CDV). The western subpopulation, which was not exposed to CDV, is a crucial element in the recovery of foxes by providing a source of animals for translocation and captive breeding. Using the program VORTEX, we developed a population viability analysis for the Santa Catalina Island fox to (1) address the likelihood of population persistence, (2) estimate the current susceptibility of the population to catastrophic events, and (3) evaluate the efficacy of current restoration strategies of releasing captive bred foxes and transplanting wild animals. Overall, we found the population to be susceptible to catastrophic events; a 50% increase in mortality every 20 years was sufficient to elevate the extinction risk above 5%. Current management activities entail the transplanting of 12 juvenile foxes annually, which may reduce the viability of the western subpopulation. A minimum population size of at least 150 foxes should be maintained in each subpopulation to reduce the risk of extinction due to demographic stochasticity. Releases of translocated and captive bred animals affect the speed of recovery on the eastern half of Catalina Island, but not the probability of extinction, which is near zero under current conditions. We conducted a sensitivity analysis for demographic parameters by incrementally varying survival, fecundity and density-dependence parameters, while holding all other parameters constant. Sensitivity analyses identified mortality and mean litter size as the most sensitive parameters, while the implementation of density-dependence and environmental variation of model parameters did not seem to affect population performance. We conclude that the population of island foxes on Santa Catalina is currently at a critically low population level, but recovery of the species appears possible.     

Optimal harvesting of ecologically interdependent fish species

The optimal exploitation of a two-species predator-prey system is considered, using Lotka-Volterra-type equations. Due to the density-dependence of ecological efficiency, both species should be harvested simultaneously over a range of relative prices. Beyond the limits of this price range, either the prey species should be utilized indirectly by harvesting the predator, or the predator should be eliminated to maximize the prey yield. Neglecting harvesting costs, the simultaneous harvest of prey and predators requires that a unit of prey biomass increase in value by being “processed” by predators. Certain results from single-species fishery models are shown not to apply to multispecies models. These are as follows: (i) Optimal regulation of a free access fishery may call for subsidizing instead of taxing the harvest of predator species. (ii) Increasing the discount rate may, at “moderate” levels, imply that the optimal standing stock of biomass increases instead of decreasing. (iii) A rising price or a falling cost per unit fishing effort of a species may raise and not lower the optimal standing stock of that species.     

Trading-off risks from predators and from aggressive hosts

Some birds exploit the aggressiveness of others that are more capable of defending their nests against predators. Usually this behaviour is selected for through improved breeding success for the timid (associate) species. The risk of predation from the more aggressive (host) species is generally rare but may select against the behaviour. The breeding success of red-breasted geese (Branta ruficollis) nesting in association with peregrine falcons (Falco peregrinus) in arctic Siberia was investigated from 1995 to 1999. Nest-predation risk for geese within colonies was positively correlated with distance from the falcons' eyries and negatively correlated with colony size. Though predation risk from the hosts was low, the risk of being attacked by the falcons was high, which we suggest explains why the likelihood of geese deserting their nests was negatively correlated with distance from the eyries. The optimal distance geese nested from eyries was 46 m, though only 37% of pairs managed to nest within 20 m of this. In years with low predation pressure, some geese within colonies nested farther from the falcons. This may be the first evidence to suggest a trade-off between costs and benefits in a nesting association, both of which are directly mediated by the host's nest defence behaviour.     

Performance of a Bayesian state-space model of semelparous species for stock-recruitment data subject to measurement error

Measurement errors in spawner abundance create problems for fish stock assessment scientists. To deal with measurement error, we develop a Bayesian state-space model for stock-recruitment data that contain measurement error in spawner abundance, process error in recruitment, and time series bias. Through extensive simulations across numerous scenarios, we compare the statistical performance of the Bayesian state-space model with that of standard regression for a traditional stock-recruitment model that only considers process error. Performance varies depending on the information content in data, as determined by stock productivity, types of harvest situations, and amount of measurement error. Overall, in terms of estimating optimal spawner abundance S_MSY, the Ricker density-dependence parameter β, and optimal harvest rate h_MSY, the Bayesian state-space model works best for informative data from low and variable harvest rate situations for high-productivity salmon stocks. The traditional stock-recruitment model (TSR) may be used for estimating α and h_MSY for low-productivity stocks from variable and high harvest rate situations. However, TSR can severely overestimate S_MSY when spawner abundance is measured with large error in low and variable harvest rate situations. We also found that there is substantial merit in using h_MSY (or benchmarks derived from it) instead of S_MSY as a management target.     

Seasonal density dependence, timing of mortality, and sustainable harvesting

Birth-pulse populations are often characterized with discrete-time models, that use a single function to relate the post-breeding population size to the post-breeding size of the previous year. Recently, models of seasonal density dependence have been constructed that emphasize interactions during shorter time periods also. Here, we study two very simple forms of density-dependent mortality, that lead to Ricker and Beverton-Holt type population dynamics when viewed over the whole year. We explore the consequences of harvest timing to equilibrium population sizes under such density dependence. Whether or not individual mortality compensates for the harvested quota, the timing of harvesting has a strong impact on the sustainability of a harvesting quota. Further, we show that careless discretization of a continuous mortality scheme may seriously underestimate the reduction in population size caused by hunting and overestimate the sustainable yield. We also introduce the concept of the demographic value of an individual, which reflects the expected contribution to population size over time in the presence of density dependence. Finally, we discuss the possibility of calculating demographic values as means of optimizing harvest strategies. Here, a Pareto optimal harvest strategy will minimize the loss of demographic value from the population for a given yield.     

Robust population management under uncertainty for structured population models.

Structured population models are increasingly used in decision making, but typically have many entries that are unknown or highly uncertain. We present an approach for the systematic analysis of the effect of uncertainties on long-term population growth or decay. Many decisions for threatened and endangered species are made with poor or no information. We can still make decisions under these circumstances in a manner that is highly defensible, even without making assumptions about the distribution of uncertainty, or limiting ourselves to discussions of single, infinitesimally small changes in the parameters. Suppose that the model (determined by the data) for the population in question predicts long-term growth. Our goal is to determine how uncertain the data can be before the model loses this property. Some uncertainties will maintain long-term growth, and some will lead to long-term decay. The uncertainties are typically structured, and can be described by several parameters. We show how to determine which parameters maintain long-term growth. We illustrate the advantages of the method by applying it to a Peregrine Falcon population. The U.S. Fish and Wildlife Service recently decided to allow minimal harvesting of Peregrine Falcons after their recent removal from the Endangered Species List. Based on published demographic rates, we find that an asymptotic growth rate lambda > 1 is guaranteed with 5% harvest rate up to 3% error in adult survival if no two-year-olds breed, and up to 11% error if all two-year-olds breed. If a population growth rate of 3% or greater is desired, the acceptable error in adult survival decreases to between 1% and 6% depending of the proportion of two-year-olds that breed. These results clearly show the interactions between uncertainties in different parameters, and suggest that a harvest decision at this stage may be premature without solid data on adult survival and the frequency of breeding by young adults.     

Cost‐Effectiveness of Translocation Options for a Threatened Waterbird

Abstract: Reintroduction of captive‐reared animals has become increasingly popular in recent decades as a conservation technique, but little is known of how demographic factors affect the success of reintroductions. We believe whether the increase in population persistence associated with reintroduction is sufficient to warrant the cost of rearing and relocating individuals should be considered as well. We examined the trade‐off between population persistence and financial cost of a reintroduction program for Crested Coots (Fulica cristata). This species was nearly extirpated from southern Europe due to unsustainable levels of hunting and reduction in amount and quality of habitat. We used a stochastic, stage‐based, single‐sex, metapopulation model with site‐specific parameters to examine the demographic effects of releasing juveniles or adults in each population for a range of durations. We parameterized the model with data from an unsuccessful reintroduction program in which juvenile captive‐bred Crested Coots were released between 2000 and 2009. Using economic data from the captive‐breeding program, we also determined whether the strategy that maximized abundance coincided with the least expensive strategy. Releasing adults resulted in slightly larger final abundance than the release of nonreproductive juveniles. Both strategies were equally poor in achieving a viable metapopulation, but releasing adults was 2–4 times more expensive than releasing juveniles. To obtain a metapopulation that would be viable for 30 years, fecundity in the wild would need to increase to the values observed in captivity and juvenile survival would need to increase to almost unity. We suggest that the most likely way to increase these vital rates is by increasing habitat quality at release sites.     

Minimise long-term loss or maximise short-term gain?: Optimal translocation strategies for threatened species

Translocation is a useful management option for conservation of threatened animal species. It can be used to increase the range of a species, augment the numbers in a critical population, or establish new populations and hence spread the risk of extinction through local catastrophes. As it is an important and expensive conservation tool, translocation management decisions must be carefully considered, with the objective of the translocation project in mind. By analysing the translocation problem within a decision-theory framework, we find optimal management decisions that are rational and transparent. We illustrate our approach using a case study of the bridled nailtail wallaby (Onychogalea fraenata). Our particular translocation question is: if we have a set number of wallabies to translocate in each time period and two translocation sites, how many wallabies should we put at each site given the state of each population to maximise the benefit to the species? We model the translocated populations with first-order Markov chain stochastic population models, and use stochastic dynamic programming to determine the optimal management decisions. We look at two sites with different growth rates – one increasing and one decreasing – and compare the optimal strategies for two different objective functions. The first is a long-term persistence objective function, which maximises the persistence of translocated populations a large number of time steps after the end of the translocation program. The second maximises total population size at the end of the translocation program. Although these objective functions are similar, they generate surprisingly different optimal translocation strategies. When maximising the long-term persistence of the translocated populations, translocation decisions are not important as long as an increasing population is established. This indicates that site quality – rather than the number and timing of translocations – primarily determines the long-term persistence of populations. When maximising total population size, the optimal strategy is to add to the increasing population unless it is above a size where it is likely to reach its carrying capacity over the planning timeframe. As translocation decisions are important in fulfilling the objective, this objective function is more useful in creating practical advice for translocation managers. The discrepancy between the optimal strategies given by the two objectives demonstrates the importance of careful consideration when specifying the goals of a project. This observation applies not only to translocation programs, but any project where clear decision-making is needed.     

Analytical Solutions to Trade‐Offs between Size of Protected Areas and Land‐Use Intensity

Abstract: Land‐use change is affecting Earth's capacity to support both wild species and a growing human population. The question is how best to manage landscapes for both species conservation and economic output. If large areas are protected to conserve species richness, then the unprotected areas must be used more intensively. Likewise, low‐intensity use leaves less area protected but may allow wild species to persist in areas that are used for market purposes. This dilemma is present in policy debates on agriculture, housing, and forestry. Our goal was to develop a theoretical model to evaluate which land‐use strategy maximizes economic output while maintaining species richness. Our theoretical model extends previous analytical models by allowing land‐use intensity on unprotected land to influence species richness in protected areas. We devised general models in which species richness (with modified species‐area curves) and economic output (a Cobb–Douglas production function) are a function of land‐use intensity and the proportion of land protected. Economic output increased as land‐use intensity and extent increased, and species richness responded to increased intensity either negatively or following the intermediate disturbance hypothesis. We solved the model analytically to identify the combination of land‐use intensity and protected area that provided the maximum amount of economic output, given a target level of species richness. The land‐use strategy that maximized economic output while maintaining species richness depended jointly on the response of species richness to land‐use intensity and protection and the effect of land use outside protected areas on species richness within protected areas. Regardless of the land‐use strategy, species richness tended to respond to changing land‐use intensity and extent in a highly nonlinear fashion.     

Should managed populations be monitored every year?

We often need to estimate the size of wild populations to determine the appropriate management action, for example, to set a harvest quota. Monitoring is usually planned under the assumption that it must be carried out at fixed intervals in time, typically annually, before the harvest quota is set. However, monitoring can be very expensive, and we should weigh the cost of monitoring against the improvement that it makes in decision making. A less costly alternative to monitoring annually is to predict the population size using a population model and information from previous surveys. In this paper, the problem of monitoring frequency is posed within a decision-theory framework. We discover that a monitoring regime that varies according to the state of the system can outperform fixed-interval monitoring. This idea is illustrated using data for a red kangaroo (Macropus rufus) population in South Australia. Whether or not one should monitor in a given year is dependent on the estimated population density in the previous year, the uncertainty in that population estimate, and past rainfall. We discover that monitoring is important when a model-based prediction of population density is very uncertain. This may occur if monitoring has not taken place for several years, or if rainfall has been above average. Monitoring is also important when prior information suggests that the population is near a critical threshold in population abundance. However, monitoring is less important when the optimal management action would not be altered by new information.     

Incorporating known sources of uncertainty to determine precautionary harvests of saltwater crocodiles.

It has been demonstrated repeatedly that the degree to which regulation operates and the magnitude of environmental variation in an exploited population will together dictate the type of sustainable harvest achievable. Yet typically, harvest models fail to incorporate uncertainty in the underlying dynamics of the target population by assuming a particular (unknown) form of endogenous control. We use a novel approach to estimate the sustainable yield of saltwater crocodile (Crocodylus porosus) populations from major river systems in the Northern Territory, Australia, as an example of a system with high uncertainty. We used multimodel inference to incorporate three levels of uncertainty in yield estimation: (1) uncertainty in the choice of the underlying model(s) used to describe population dynamics, (2) the error associated with the precision and bias of model parameter estimation, and (3) environmental fluctuation (process error). We demonstrate varying strength of evidence for density regulation (1.3-96.7%) for crocodiles among 19 river systems by applying a continuum of five dynamical models (density-independent with and without drift and three alternative density-dependent models) to time series of density estimates. Evidence for density dependence increased with the number of yearly transitions over which each river system was monitored. Deterministic proportional maximum sustainable yield (PMSY) models varied widely among river systems (0.042-0.611), and there was strong evidence for an increasing PMSY as support for density dependence rose. However, there was also a large discrepancy between PMSY values and those produced by the full stochastic simulation projection incorporating all forms of uncertainty, which can be explained by the contribution of process error to estimates of sustainable harvest. We also determined that a fixed-quota harvest strategy (up to 0.2K, where K is the carrying capacity) reduces population size much more rapidly than proportional harvest (the latter strategy requiring temporal monitoring of population size to adjust harvest quotas) and greatly inflates the risk of resource depletion. Using an iconic species recovering from recent extreme overexploitation to examine the potential for renewed sustainable harvest, we have demonstrated that incorporating major forms of uncertainty into a single quantitative framework provides a robust approach to modeling the dynamics of exploited populations.     

Uncertainty in age-specific harvest estimates and consequences for white-tailed deer management

Age structure proportions (proportion of harvested individuals within each age class) are commonly used as support for regulatory restrictions and input for deer population models. Such use requires critical evaluation when harvest regulations force hunters to selectively harvest specific age classes, due to impact on the underlying population age structure. We used a stochastic population simulation model to evaluate the impact of using harvest proportions to evaluate changes in population age structure under a selective harvest management program at two scales. Using harvest proportions to parameterize the age-specific harvest segment of the model for the local scale showed that predictions of post-harvest age structure did not vary dependent upon whether selective harvest criteria were in use or not. At the county scale, yearling frequency in the post-harvest population increased, but model predictions indicated that post-harvest population size of 2.5 years old males would decline below levels found before implementation of the antler restriction, reducing the number of individuals recruited into older age classes. Across the range of age-specific harvest rates modeled, our simulation predicted that underestimation of age-specific harvest rates has considerable influence on predictions of post-harvest population age structure. We found that the consequence of uncertainty in harvest rates corresponds to uncertainty in predictions of residual population structure, and this correspondence is proportional to scale. Our simulations also indicate that regardless of use of harvest proportions or harvest rates, at either the local or county scale the modeled SHC had a high probability (>0.60 and >0.75, respectively) of eliminating recruitment into >2.5 years old age classes. Although frequently used to increase population age structure, our modeling indicated that selective harvest criteria can decrease or eliminate the number of white-tailed deer recruited into older age classes. Thus, we suggest that using harvest proportions for management planning and evaluation should be viewed with caution. In addition, we recommend that managers focus more attention on estimation of age-specific harvest rates, and modeling approaches which combine harvest rates with information from harvested individuals to further increase their ability to effectively manage deer populations under selective harvest programs.