Publications

Publications & Presentations

Video Lectures

Re-interpreting the Fisheries Crisis

In Press

Minte-Vera, C.V., R. Hilborn. Bayesian hierarchical meta-analysis of density-dependent body growth in haddock (Melanogrammus aeglefinus). Can. J. Fish. Aquat. Sci.

Abstract

The evidence for changes in somatic growth related to population density was investigated for eleven stocks of haddock using Bayesian hierarchic meta-analysis. A within-stock growth model that includes density-dependence in the growth increment of ages 2 and older was used. The posterior distribution for the hyper-mean of the density-dependence parameter indicated overall evidence of negative density-dependence, owever, there was considerable heterogeneity among stocks. Five stocks had negative density-dependence (Georges Bank, Eastern Scotian Shelf, North East Artic, West of Scotland and Iceland), three stocks had positive density-dependence (Bay of Fundy/Scotian Shelf, North Sea/Skagerrak and West of Scotland), one stock had density-dependence around zero (Faroes), the remainders had little information about density-dependent on their data (Irish Sea and Celtic Sea/West of Ireland) and thus their estimate was closer to the prediction from the hierarchical model for an unobserved stock, which includes the information for all the stocks.

In Review

Scheuerell, M.D., R. Hilborn. Estimating the freshwater component of essential fish habitat for Pacific salmon (Oncorhynchus spp.) with the Shiraz model.

Abstract

The 1996 Sustainable Fisheries Act contained provisions that all federal fisheries management plans should contain some description of essential fish habitat (EFH). While much emphasis has been placed on estimating EFH for marine stocks, very little attention has been paid to doing so for Pacific salmon, in part due to their complex life history strategies. An earlier assessment of EFH for Pacific salmon across the west coast of the United State focused on the freshwater component of EFH due to limited knowledge about marine distributions. That analysis concluded that a more in-depth examination of how freshwater habitat affects the various life stages, at much smaller spatial scales than previously considered, was indeed necessary. Here we used a detailed life history model for Pacific salmon to estimate the freshwater component of EFH for two populations of Chinook salmon listed as threatened under the Endangered Species Act within a large watershed draining into Puget Sound, Washington, U.S.A. By accounting for proposed harvest rates, hatchery practices, and habitat structure, we were able to identify 23 of 60 subbasins that were essential fish habitat for insuring that there would be no significant decline in the total number of spawners relative to the current average escapement. Our analytical framework could be easily applied to other populations or species of salmon to aid in developing recovery and management plans.

Hilborn, R. Life history models for salmon management: the challenges.

Abstract

Salmon have complex life histories that have been extensively studied, particularly in freshwater, yet most salmon management relies on models that ignore much of salmon life history. For instance, calculation of optimal escapement for most Pacific salmon stocks summarizes their entire life history into a single relationship between spawners and subsequent recruits. Similarly, most analyses of salmon habitat have used models that fail to integrate the complex life history of salmon and have often considered only a single “limiting factor”. Computational methods and models are now being used to incorporate life history and habitat information directly into evaluations of both harvesting and habitat management policies. Challenges and opportunities in using life history models include (1) the need for better dynamic understanding of how habitat affects survival, (2) turning current “expert system” analysis into statistical estimation, (3) application of life history models to hatchery/wild interaction, (4) quantifying essential fish habitat using life history models and (5) use of such models to explore salmon/ocean interactions.

Magnusson, A., R. Hilborn. Information about fish stock abundance: Data, models, and assumptions.

Abstract

Informative data in fisheries stock assessment are those that lead to accurate estimates of abundance and reference points. In practice, the accuracy of estimated abundance is unknown and it’s often unclear which features of the data make them informative or uninformative. Neither is it obvious which model assumptions will improve estimation performance, given a particular dataset. In this simulation study, 10 hypotheses are addressed using multiple scenarios, estimation models, and reference points. The simulated data scenarios all share the same biological and fleet characteristics, but vary in terms of the fishing history. The estimation models are based on a common statistical catch-at-age framework, but estimate different parameters and have different parts of the data available to them. Among the findings is that a “one-way trip” scenario, where harvest rate gradually increases while abundance decreases, proved no less informative than a contrasted catch history. Models that excluded either abundance index or catch at age performed surprisingly well, compared to models that included both data types. Natural mortality rate, M, was estimated with some reliability when age composition data were available from before major catches were removed. Stock-recruitment steepness, h, was estimated with some reliability when abundance index or age composition data were available from years of very low abundance. Understanding what makes fisheries data informative or uninformative enables scientists to identify fisheries for which stock assessment models are likely to be biased or imprecise. Managers can also benefit from guidelines on how to distribute funding and manpower among different data collection programmes to gather the most information.

Hilborn, R., C. Minte-Vera. Fisheries induced changes in growth rates in marine fisheries: are they significant?

Abstract

Fishing provides selective pressure on many fisheries life history traits, and there has long been an interest in the impact of size selective fishing on the evolution of growth rates. Recent studies, both laboratory and empirical, suggest that such size selective fishing may be significant. Using a meta analysis of 73 commercially fished stocks we show that declines in mass-at-age are slightly more common than increases, but there is no relationship between the intensity of fishing and the change in growth rate. We review a number of size selectivity patterns in major commercial fisheries and show that the intensity of selection and the size selectivity are both considerably less than used in laboratory experiments. We simulate the evolutionary impact of fishing on growth and show that given the actual selectivity patterns found in most commercial fisheries little evolutionary impact in growth rates is expected. The model shows the best way to reduce evolutionary impacts is to lower exploitation rates. We suggest that for fisheries where there is very intense size specific selection, managers would be advised to use a model such as ours to evaluate potential evolutionary impacts.

Lessard, R.B., R. Hilborn, B. Chasco. Beyond brood tables: life-history models of sockeye salmon (Oncorynchus nerka) populations.

Abstract

We compare two methods of analyzing stock recruitment relationships. Our primary concern is to develop a new method to establish harvest goals for sockeye salmon populations in Bristol Bay Alaska. We model the life history of a fish population from a spawning stage, through juvenile and adult stages, and ending with adults that return to spawn. We specifically model a Bristol Bay sockeye population composed of age-classes coming from 1 or 2 years of lake rearing and 2 or 3 years of ocean residency. We initially place density dependence in the egg to fry stage. We fit the model to spawner and recruit data with each adult age class represented and compare the results of fitting the model with and without smolt composition data. Parameters are estimated by maximizing a likelihood objective function. Posterior estimates of parameters from MCMC simulations are then used to assess optimal harvest policies. We search for policies that produce the highest average yield under the assumption that estimated parameters reflect the best estimate of the ”state-of-nature” of the system. We found that it is possible to detect density dependence with a life history model where analysis of Beverton-Holt stock recruitment relationship fails to do so. We find that Beverton-Holt relationships produce policies and long term yield estimates that are inconsistent with empirical trends. Conversely, we find that harvest rates and maximum sustained yield estimates using the life history model estimate are consistent with the historical behavior of fisheries examined.

PDFs

Please contact rayh@u.washington.edu to obtain PDF copies. Links to PDFs (Acrobat Reader required) of various Hilborn publications will be provided when possible.

Citations

Books and Monographs

Punt, A. and R. Hilborn. 2002. Bayesian stock assessment methods in fisheries. FAO Computerized Information Series (Fisheries) No. 12. 56 p.

Hilborn, R. and M. Mangel. 1997. The Ecological Detective: confronting models with data. Princeton University Press, Princeton, N.J. 315 pps.

Punt, A.E. and R. Hilborn. 1996. Biomass dynamics models. FAO Computerized Information Series (Fisheries). No. 10. Rome, FAO. 62p.

Hilborn, R. and C. J. Walters. 1992. Quantitative Fisheries Stock Assessment: Choice, Dynamics and Uncertainty. Chapman and Hall, New York. 570 p. Also available in Russian.

Bazykin, A., P. Bunnell, W.C. Clark, G.C. Gallopin, J. Gross, R. Hilborn, C.S. Holling, D.D. Jones, R.M. Peterman, J.E. Rabinovich, J.H. Steele, and C.J. Walters. 1978. Adaptive Environmental Assessment and Management. John Wiley and Sons, New York. 375 pps.

Refereed Journals

Bue, B.G., R. Hilborn, and M.R. Link. 2008. Optimal harvesting considering biological and economic objectives. Can. J. Fish. Aquat. Sci. 65:691-700.

Most examinations of optimal harvesting policies have considered only biological objectives, yet it is increasingly recognized that a primary objective of many fisheries is economic profitability. Using Bayesian risk analysis, we compare policies that combine fisheries harvesting, the revenue brought in by fish sales, the cost of harvesting and processing, and processing and fishing capacity to find policies that maximize biological yield and economic profit to the processing and harvesting sectors, for a major Pacific salmon (Oncorhynchus spp.) fishery in Bristol Bay, Alaska. We show that while average catch is maximized by a fixed escapement policy, total revenue is maximized by a policy that includes some harvesting at stock sizes below that required to produce maximum average catch. In addition, there is a wide range of policies that provide 90% of the maximum for any of the biological and economic objectives considered and economic profitability is enhanced by limitations on processing and harvesting capacity.

Grafton, RQ, R Hilborn, L Ridgeway, D Squires, M Williams, S Garcia, T Groves, J Joseph, K Kelleher, T Kompas, G Libecap, CG Lundin, M Makino, T Matthiasson, R McLoughlin, A Parma, G San Martin, B Satia, C-C Schmidt, M Tait, LX Zhang. 2008. Positioning fisheries in a changing world. Mar. Pol. 32:630-634.

Marine capture fisheries face major and complex challenges: habitat degradation, poor economic returns, social hardships from depleted stocks, illegal fishing, and climate change, among others. The key factors that prevent the transition to sustainable fisheries are information failures, transition costs, use and non-use conflicts and capacity constraints. Using the experiences of fisheries successes and failures it is argued only through better governance and institutional change that encompasses the public good of the oceans (biodiversity, ecosystem integrity, sustainability) and societal values (existence, aesthetic and amenity) will fisheries be made sustainable.

Gunderson, DR, AM Parma, R Hilborn, JM Cope, DL Fluharty, ML Miller, RD Vetter, SS Heppell, HG Greene. 2008. The challenge of managing nearshore rocky reef resources. Fisheries 33(4):172-179.

Nearshore temperate reefs are highly diverse and productive habitats that provide structure and shelter for a wide variety of fishes and invertebrates. Recreational and commercial fisheries depend on nearshore reefs, which also provide opportunities for non-extractive recreational activities such as diving. Many inhabitants of nearshore temperate reefs on the west coast of North America have very limited home ranges as adults, and recent genetic evidence indicates that the dispersion of the larval stages is often restricted to tens of kilometers. Management of temperate reef resources must be organized on very small spatial scales in order to be effective, offering unique technical challenges in terms of assessment and monitoring. New enabling legislation could assist in specifying mandates and adjusting institutional design to allow stakeholders and concerned citizens to formulate management policies at local levels, and to aid in implementing and enforcing these policies.

Hard, JJ, MR Gross, M Heino, R Hilborn, RG Kope, R Law, JD Reynolds. 2008. Evolutionary consequences of fishing and their implications for salmon. Evol. Applic. 1:388-408. DOI: 10.1111/j.1752-4571.2008.00020.x

We review the evidence for fisheries-induced evolution in anadromous salmonids. Salmon are exposed to a variety of fishing gears and intensities as immature or maturing individuals. We evaluate the evidence that fishing is causing evolutionary changes to traits including body size, migration timing and age of maturation, and we discuss the implications for fisheries and conservation. Few studies have fully evaluated the ingredients of fisheries-induced evolution: selection intensity, genetic variability, correlation among traits under selection, and response to selection. Most studies are limited in their ability to separate genetic responses from phenotypic plasticity, and environmental change complicates interpretation. However, strong evidence for selection intensity and for genetic variability in salmon fitness traits indicates that fishing can cause detectable evolution within ten or fewer generations. Evolutionary issues are therefore meaningful considerations in salmon fishery management. Evolutionary biologists have rarely been involved in the development of salmon fishing policy, yet evolutionary biology is relevant to the long-term success of fisheries. Future management might consider fishing policy to (i) allow experimental testing of evolutionary responses to exploitation and (ii) improve the long-term sustainability of the fishery by mitigating unfavorable evolutionary responses to fishing. We provide suggestions for how this might be done.

Hilborn, R. 2008. Knowledge on how to achieve sustainable fisheries. Pages 45-56 in K Tsukamoto, T Kawamura, T Takeuchi, TD Beard, Jr, MJ Kaiser (eds), Fisheries for Global Welfare and Environment, 5th World Fisheries Congress 2008.

I review the state of current knowledge with respect to the requirements for achieving sustainable fisheries. I consider the range of objectives for fisheries and identify conflicting objectives as a major issue in achieving sustainability. Next I review historical and current practice in allocation of fish resources and regulation of harvest and highlight existing knowledge. Evidence suggests that both restriction of access and maintenance of biological productivity are necessary conditions to achieve biological, economic and social sustainability. However, the tools appropriate to achieve these differ greatly across fisheries and societies, and for both elements of fisheries management localsolutions are needed in most cases. Attempts to impose standardized solutions to either issue frequently result in ineffective solutions. Evidence also suggests that involvement of consumptive users through appropriate incentives is an essential element in achieving sustainability.

Hilborn, R, CV Minte-Vera. 2008. Fisheries-induced changes in growth rates in marine fisheries: are they significant? Bull. Mar. Sci. 83(1):95-105.

Fishing provides selective pressure on many fisheries life-history traits, and interest in the impact of size-selective fishing on the evolution of growth rates is long standing. Recent studies, both laboratory and empirical, suggest that such size-selective fishing is significant. Using a metaanalysis of 73 commercially fished stocks, we found that declines in mass at age are slightly more common than increases, but no relationship was apparent between the intensity of fishing and the change in growth rate. We reviewed a number of size-selectivity patterns in major commercial fisheries and found that the intensity of selection and the size selectivity were both considerably less than are used in laboratory experiments. We simulated the evolutionary impact of fishing on growth and found that, given the actual selectivity patterns found in most commercial fisheries, little evolutionary impact on growth rates is expected. The model showed that the best way to reduce evolutionary impacts is to lower exploitation rates. We suggest that, for fisheries where size-specific selection is very intense, managers should use a model such as ours to evaluate potential evolutionary impacts.

Lessard RB, R Hilborn, BE Chasco. 2008. Escapement goal analysis and stock reconstruction of sockeye salmon populations (Oncorhynchus nerka) using life-history models. Can. J. Fish. Aquat. Sci. 65(10):2269–2278. Available online.

We compare life-history models with the Beverton–Holt approach of escapement goal analysis. We model the life history of a sockeye salmon (Onchorhynchus nerka) population from a spawning stage, through juvenile and adult stages, and ending with adults that return to spawn. We fit models to data by statistically comparing predicted and observed numbers of four dominant adult ages. Posterior estimates of parameters from Markov chain Monte Carlo simulations are then used to assess optimal harvest policies. We search for policies that produce the highest average yield. We find that it is possible to detect density dependence with a life-history model where analysis of Beverton–Holt stock–recruitment relationship fails to do so. We find that Beverton–Holt relationships produce policies and long-term yield estimates that are inconsistent with empirical trends. Conversely, we find that optimal spawning stock sizes and maximum sustained yield estimates using the life-history model estimate are consistent with the historical behavior of fisheries examined. Adding smolt data to the analysis does not substantially change predicted optimal spawning stock size, but decreases the variance in estimated posterior parameter distributions and policy variable distributions.

Lin, J, TP Quinn, R Hilborn, L Hauser. 2008. Fine-scale differentiation between sockeye salmon ecotypes and the effect of phenotype on straying. Heredity 101:341-350.

A long-standing goal of evolutionary biology is to understand the factors that drive population divergence and speciation, and conspecific ecotypes are considered an intermediate step towards speciation. Competing hypotheses for differentiation among salmonid ecotypes include geographic separation, accurate homing, and selection against introgression. Here, we used genetic and phenotypic data from geographically proximate populations of beach and stream ecotypes of sockeye salmon (Oncorhynchus nerka) in Little Togiak Lake, Alaska, to examine the relationship between ecological and genetic differentiation. Both genetic and phenotypic differentiation was high and significant between beach and creek samples in all years. Within ecotypes, beach spawners showed lower differentiation (FST =0.007) and higher genetic variability (HE=0.789) than stream spawners (FST=0.038, HE =0.730). Fish genetically identified as strays differed phenotypically from their resident conspecifics: males collected in a creek but genetically assigned to the beach population had shallower body depths (similar to native creek fish) than males assigned to and sampled on beaches, and males genetically assigned to creeks but collected on a beach were deeper-bodied than males genetically assigned to and sampled in the creeks. Thus the fish that strayed tended to resemble the morphology of the fish in the population that they joined.

Martell, S.J.D., C.J. Walters, and R. Hilborn. 2008. Retrospective analysis of harvest management performance for Bristol Bay and Fraser River sockeye salmon (Oncorhynchus nerka). Can. J. Fish. Aquat. Sci. 65:409-424.

Given current knowledge of mean stock-recruitment relationships and past recruitment anomalies due to environmental factors, and absent constraints on exploitation rates due to mixed stock harvesting, yield of sockeye salmon in Bristol Bay Alaska and Fraser River B.C. might have been at least 100% larger since 1950 than was actually achieved, and possibly as much as 300% larger depending on responses of a few large stocks for which the optimum stock size remains highly uncertain. Most of these gains would have been due to knowledge of optimum mean spawning stock size rather than specific recruitment anomalies; knowing all future recruitment anomalies at the time of each spawning stock choice would have likely only added 2-5% to total catches. For some stocks, delayed density dependence (cyclic dominance) might have resulted in somewhat lower yields, but under optimal management would still have been higher than were achieved. Even given only estimates of optimum spawning stock size each year based on data available as of that year, but following fixed escapement harvest policy rules, managers could likely have achieved 30-40% higher total yield. Key management experiments for the future will involve testing for cyclic dominance effects on two major stocks (Kvichak, Late Shuswap), to determine whether stocks with strong delayed density dependent survival effects should be deliberately managed through fallow rotation strategies for juvenile nursery lakes.

de Mutsert, K, JH Cowan, Jr, TE Essington, R Hilborn. 2008. Reanalyses of Gulf of Mexico fisheries data: Landings can be misleading in assessments of fisheries and fisheries ecosystems. PNAS 105(7):2740-2744.

We used two high profile articles as cases to demonstrate that use of fishery landings data can lead to faulty interpretations about the condition of fishery ecosystems. One case uses the mean trophic level index and its changes, and the other uses estimates of fishery collapses. In earlier analyses by other authors, marine ecosystems in the Gulf of Mexico (GOM) and U.S. Atlantic Ocean south of Chesapeake Bay were deemed to be severely overfished and the food webs badly deteriorated using these criteria. In our reanalyses, the low mean trophic level index for the GOM actually resulted from large catches of two groups of low trophic level species, menhaden and shrimp, and the mean trophic level was slowly increasing rather than decreasing. Commercial targeting and high landings of shrimps and menhaden, especially in the GOM, drove the index as previously calculated. Reanalyses of fishery collapses incorporating criteria that included targeting, variability in fishing effort, and market forces discovered many false cases of collapse based simply upon a decline of catches to 10% of previous maximum levels. Consequently, we suggest that the low mean trophic level index calculated in the earlier article for the GOM did not reflect the overall condition of the fishery ecosystem, and that the 10% rule for collapse should not be interpreted out of context in the GOM or elsewhere. In both cases, problems lay in the assumption that commercial landings data alone adequately reflect the fish populations and communities.

Sethi, S.A., and R. Hilborn. 2008. Interactions between poaching and management policy affect marine reserves as conservation tools. Biol. Cons. 141:506-516.

This analysis uses a simple age-structured reserve model with Black rockfish biology to explore the effects of poaching within reserve boundaries under three different management policies based on yield maximization or reproductive thresholds. Departures from the traditional assumptions of full compliance to reserve boundaries alter the conclusions of prior modeling work that demonstrate yield equivalence to no-reserve effort control management and augmented reproductive benefits when small reserves are implemented. By degrading the recruitment subsidization effect to nonreserve areas from protected reserve populations, poaching resulted in negative externalities for compliant fishermen in open areas in terms of yield and degraded the reproductive output and age-structure of the system. All three policies required effort reduction in open areas as a response to poaching in reserves. The strength of the impacts from poaching varied with policy choice and harvest intensity in the reserve, where at the highest level of poaching modeled here (15% annual exploitation rate of the vulnerable reserve population) biological and fishery benefits of implementing reserves were totally negated. Under the assumptions of this model, a policy managing for a reproductive threshold that excludes the reserve population is the precautionary choice if poaching is likely. The results of this exercise emphasize the importance of garnering compliance to reserve boundaries from resource-users for spatial closures to be successful ocean management tools.

Westley, PAH, R Hilborn, TP Quinn, GT Ruggerone, DE Schindler. 2008. Long-term changes in rearing habitat and downstream movement by juvenile sockeye salmon (Oncorhynchus nerka) in an interconnected Alaska lake system. Ecol. Freshwat. Fish. 17:443–454.

In some populations the phenomenon of partial migration develops where some individuals stay in a given habitat rather than move with the migratory component. Depending on the selective pressures, the individuals that stay may be larger, smaller or similar in size to those that move. Freshwater movements of juvenile sockeye salmon (Oncorhynchus nerka Walbaum) fry vary among and within populations, and can be complex, especially in interconnected lake systems. We examined variation of movement patterns by a sockeye salmon population in an interconnected lake system during a period of rapid natural habitat change and found that fry migrating downstream were shorter, had lower body condition, and were more likely ill and moribund compared with fish remaining in the lake. However, otolith microstructure measurements indicated that emigrants did not grow significantly slower than residents prior to downstream movement. We show that patterns (i.e., demography of migrants, timing of movement) of downstream movement have changed since the 1970s, corresponding to changes in rearing habitat. Our findings parallel the results with other salmonid species and are generally consistent with the paradigm that density-dependent interactions from declining habitat availability or quality result in the downstream movement of competitively inferior individuals, although the mechanisms governing downstream migration are unclear in this system

Carlson, S.M., R. Hilborn, A.P. Hendry, and T.P. Quinn. 2007. Predation by bears drives senescence in natural populations of salmon. PLoS One 2(12):ed1286. doi:10.1371/journal.pone.0001286.

Classic evolutionary theory predicts that populations experiencing higher rates of environmentally caused (“extrinsic”) mortality should senesce more rapidly, but this theory usually neglects plausible relationships between an individual's senescent condition and its susceptibility to extrinsic mortality. We tested for the evolutionary importance of this condition dependence by comparing senescence rates among natural populations of sockeye salmon (Oncorhynchus nerka) subject to varying degrees of predation by brown bears (Ursus arctos). We related senescence rates in six populations to (1) the overall rate of extrinsic mortality, and (2) the degree of condition dependence in this mortality. Senescence rates were determined by modeling the mortality of individually-tagged breeding salmon at each site. The overall rate of extrinsic mortality was estimated as the long-term average of the annual percentage of salmon killed by bears. The degree of condition dependence was estimated as the extent to which bears killed salmon that exhibited varying degrees of senescence. We found that the degree of condition dependence in extrinsic mortality was very important in driving senescence: populations where bears selectively killed fish showing advanced senescence were those that senesced least rapidly. The overall rate of extrinsic mortality also contributed to among-population variation in senescence-but to a lesser extent. Condition-dependent susceptibility to extrinsic mortality should be incorporated more often into theoretical models and should be explicitly tested in natural populations.

Chasco, B., R. Hilborn, and A.E. Punt. 2007. Run reconstruction of mixed-stock salmon fisheries using age-composition data. Can. J. Fish. Aquat. Sci. 64:1479-1490.

A method for using age composition data to determine stock-specific migration timing and abundance in a mixed-stock salmon fishery is developed. The Chignik sockeye fishery has two stocks, but only aggregate catch and escapement data are available. The age composition of the two stocks, however, is known to be consistently different, and age composition data are collected from one stock at the beginning of the commercial fishing season and from the commercial catch throughout the season. Using the changes in age composition in the commercial catch throughout the season we estimate the total abundance and migration timing for the two Chignik stocks using maximum likelihood and Bayesian analyses. The accuracy of this stock separation model was highly correlated with that of scale pattern analysis for most years from 1978 to 2002 (r=0.89). The results suggest that age-composition may provide salmon managers with a reliable and inexpensive method for determining stock-specific migration timing and abundance in a mixed-stock fishery.

Grafton, R.Q., T. Kompas, and R.W. Hilborn. 2007. Economics of overexploitation revisited. Science 318:1601.

Hilborn, R. 2007. Biodiversity loss in the oceans: how bad is it? Science 316:1281-1282.

Hilborn, R. 2007. Defining success in fisheries and conflicts in objectives. Marine Policy 31:153-158.

Hilborn, R. 2007. Faith, evolution, and the burden of proof—the author responds. Fisheries 32:91-93.

Hilborn, R. 2007. Managing fisheries is managing people: what has been learned? Fish and Fisheries 8:285–296.

Understanding the behaviour of fishermen is a key ingredient to successful fisheries management. The aggregate behaviour of fishing fleets can be predicted and managed with appropriate incentives. To determine appropriate incentives, we should look to successes to learn what works and what does not. In different fisheries incentive systems have been found to reduce the race-for-fish and make fisheries profitable, to stimulate stock rebuilding, to reduce bycatch, and to provide for reductions in illegal fishing. Yet, success can be evaluated in many dimensions, but is, in fact, rarely done – per cent overfished seems to be the dominant measure of performance. I evaluate the yield lost due to overfishing in several ecosystems and contrast the situation of North Atlantic cod where considerable yield is lost, to fisheries in New Zealand and the west coast of the USA where lost yield due to overfishing is very small. Much more systematic evaluation of the other aspects of fisheries performance is greatly needed. From examples explored in this paper I conclude that prevention of overfishing can be achieved with strong central governments enforcing conservative catch regulations, but economic success appears to require an appropriate incentive structure.

Hilborn, R. 2007. Moving to sustainability by learning from successful fisheries. Ambio 36(4):296-303.

There are two diverging views of the status and future of the world’s fisheries. One group represented largely by academic marine ecologists sees almost universal failure of fisheries management and calls for the use of marineprotected areas as the central tool of a new approach to rebuilding the marine ecosystems of the world. The scientists working in fisheries agencies and many academic scientists see a more complex picture, with many failed fisheries but also numerous successes. This group argues that we need to apply the lessons from the successful fisheries to stop the decline and rebuild those fisheries threatened by excess fishing. These lessons are stopping the competitive race to fish by appropriate incentives for fishing fleets and good governance. The major tool of resetting incentives is granting various forms of dedicated access, including community-based fishing rights, allocation to cooperatives, and individual fishing quotas. Many of the failed fisheries of the world occur in jurisdictions where central governments are not functional, and local control of fisheries is an essential part of the solution.

Hilborn, R. 2007. Reinterpreting the state of fisheries and their management. Ecosystems. DOI: 10.1007/s10021-007-9100-5.

Hilborn, R. 2007. Review: Return to the River: Restoring Salmon to the Columbia River, RN Williams (ed). 2006. Elsevier Academic Press, Amsterdam. Restoration Ecology 15(4):747–748.

Hilborn, R, G Hopcraft, P Arcese. 2007. Wildlife population increases in Serengeti National Park—response. Science 315:1790-1791.

Magnusson, A., R. Hilborn. 2007. What makes fisheries data informative? Fish and Fisheries 8:337-358.

Informative data in fisheries stock assessment are those that lead to accurate estimates of abundance and reference points. In practice, the accuracy of estimated abundance is unknown and it is often unclear which features of the data make them informative or uninformative. Neither is it obvious which model assumptions will improve estimation performance, given a particular data set. In this simulation study, 10 hypotheses are addressed using multiple scenarios, estimation models, and reference points. The simulated data scenarios all share the same biological and fleet characteristics, but vary in terms of the fishing history. The estimation models are based on a common statistical catch-at-age framework, but estimate different parameters and have different parts of the data available to them. Among the findings is that a ‘one-way trip’ scenario, where harvest rate gradually increases while abundance decreases, proved no less informative than a contrasted catch history. Models that excluded either abundance index or catch at age performed surprisingly well, compared to models that included both data types. Natural mortality rate, M, was estimated with some reliability when age-composition data were available from before major catches were removed. Stock-recruitment steepness, h, was estimated with some reliability when abundance-index or age-composition data were available from years of very low abundance. Understanding what makes fisheries data informative or uninformative enables scientists to identify fisheries for which stock assessment models are likely to be biased or imprecise. Managers can also benefit from guidelines on how to distribute funding and manpower among different data collection programmes to gather the most information.

Metzger, KL, ARE Sinclair, KLI Campbell, R Hilborn, JGC Hopcraft, SAR Mduma, RM Reich. 2007. Using historical data to establish baselines for conservation: The black rhinoceros (Diceros bicornis) of the Serengeti as a case study. Biol. Cons. 139:358-374.

Using historical animal counts, human population censuses and arrest records we determined the potential contemporary distribution of the black rhino in the Serengeti National Park. Prior to extensive poaching of the black rhino in the Serengeti (1977-78), 31 monthly reconnaissance surveys (1969-72) were made over the ecosystem, recording the number and location of animals. Using these data, we determined a reliable historical population estimate for the black rhino. We also developed a habitat suitability model of the black rhino in the Serengeti National Park using the spatial location of historical count data and contemporary vegetation and landscape variables. However because illegal hunting still remains a significant threat to the persistence of the rhino, we also determined areas where the likelihood of encountering people is high. From this analysis, we determined possible locations within the park for reintroduction of the black rhino under current conditions.

Naish, KA, JE Taylor, PS Levin, TP Quinn, JR Winton, D Huppert, R Hilborn. 2007. An evaluation of the effects of conservation and fishery enhancement hatcheries on wild populations of salmon. Advanc. Mar. Biol. 53:61-194. doi:10.1016/S0065-2881(07)53002-6.

The historical, political and scientific aspects of salmon hatchery programmes designed to enhance fishery production or to sustain or recover endangered populations are reviewed. Recognizing that the establishment of hatcheries is a political response to societal demands for harvest and conservation, we critically examine the levels of activity, the biological risks, and the economic analysis associated with salmon hatchery programmes within this social context. However, a rigorous analysis of the impacts of hatchery programmes was hindered by the lack of standardized data on release sizes and survival rates at all ecological scales, and since hatchery programme objectives are rarely defined, it was also difficult to measure their effectiveness at meeting release objectives. We examined in detail the genetic and competitive outcomes, and the risks of disease transmission of releasing hatchery fish on wild populations, and the effects of harvesting mixed stocks comprising both groups. Debates on the genetic effects of hatchery programmes are dominated by whether correct management practices can reduce negative outcomes, but there is an absence of programmatic research approaches addressing this important question. The outcome of competition between hatchery and wild fish is complex and fishery enhancement programmes should seek to reduce interactions between hatchery and wild fish at all ecological scales during their life history; but these issues are rarely studied and thus are not typically considered. Recently, managers have recognized that fishing effort on salmon released from fishery enhancement hatcheries likely impacts vulnerable wild populations and have responded by mass marking hatchery fish, so that fishing effort can be directed towards hatchery populations. However, the effectiveness of the approach is dependant on accurate marking and production of hatchery fish with high survival rates, and it is not yet clear whether selective fishing on hatchery stocks will be effective. Research demonstrating disease transmission from hatchery fish to wild populations is equivocal; evidence in this area is constrained by the lack of effective approaches to studying the fate of pathogens in the wild. We review several approaches to studying the economic consequences of hatchery programmes, but recognize that placing monetary value on conservation efforts or on hatcheries that mitigate cultural groups’ loss of historical harvest opportunities is difficult. We end by identifying existing major knowledge gaps, which, if filled, could contribute towards a fuller understanding of the role that hatchery programmes could play in meeting divergent goals. However, we 5 recognize that many management recommendations arising from such research may involve trade-offs between different risks, and that decisions about these trade-offs must occur within a social context. Hatcheries have played an important role in sustaining some highly endangered populations, and it is possible that reform of conservation hatchery practices will lead to an increase in the number of successful programmes. However, a serious appraisal of the role of hatcheries in meeting broader needs, such as harvest augmentation and mitigation, is urgently warranted and should take place at the scientific, but more effectively, at the societal level.

Quinn, TP, S Hodgson, L Flynn, R Hilborn, DE Rogers. 2007. Directional selection by fisheries and the timing of sockeye salmon migrations. Ecol. Applic. 17:731-739.

Sinclair, ARE, SAR Mduma, GC Hopcraft, JM Fryxell, R Hilborn, S Thirgood. 2007. Long-term ecosystem dynamics in the Serengeti: lessons for conservation. Cons. Biol. 21(3):580-590.

Long-term ecological studies are important to understanding ecosystem dynamics and for guiding evidence-based management. In the Serengeti-Mara Ecosystem, we examined natural and anthropogenic disturbances to further understanding of how the system functions. Through long-term monitoring of different components of the system we traced the effects of disturbances to elucidate cause and effect connections between them. Our quasi-natural experiment showed how different components of the ecosystem are integrated. Long-term data illustrated the role of population regulation in mammals, particularly in migratory wildebeest and non-migratory buffalo, through food limitation. Predation limited populations of smaller resident ungulates and small carnivores. Abiotic events, such as droughts and floods, created disturbances that affected survivorship of ungulates and birds. Such disturbances showed feedbacks between the system’s biotic and abiotic realms. Interactions between elephants and their food allowed savanna and grassland communities to co-occur as multiple states. Predators made use of the increase in woodland vegetation to facilitate capture of prey. This was a non-linear indirect interaction. Anthropogenic disturbances were direct through and hunting and indirect through transfer of disease to wildlife. Slow and rapid change and multiple ecosystem states became apparent only over a period of several decades and involved events at different spatial scales.

Conservation efforts need to accommodate both infrequent and unpredictable events and long-term trends. Management should plan on the time scale of those events and should not aim to maintain the status quo. Systems can be self-regulating by either food or natural enemies; thus, culling may not be required. Conservation efforts should consider that the ecosystem can occur in multiple states, and that there is no a priori need to maintain only one natural state. Finally, conservation efforts outside protected areas must distinguish between natural change and direct human-induced change. Protected areas can act as ecological baselines where human-induced change is kept to a minimum.

Walters, CJ, R Hilborn, R Parrish. 2007. An equilibrium model for predicting the efficacy of marine protected areas in coastal environments. Can. J. Fish. Aquat. Sci. 64:1009-1018.

Quantitative models of marine protected area proposals can be used to compare outcomes given current biological and economic knowledge. We used a model of a linear coastline, broken into 200 discrete cells each spanning 1.6 km of coast. This model is used to evaluate alternative proposals for marine protected area networks, predicting long-term (equilibrium) changes in abundances and harvests while accounting for dispersal of both larvae and older fish, changes in mean fecundity with reduced mortality in reserves, impacts of displaced fishing effort on abundances outside reserves, and compensatory (stock-recruitment) changes in post-settlement juvenile survival. The model demonstrates that even modest dispersal rates of older fish can substantially reduce the increase of abundance within protected areas compared to predictions from simpler models that ignore such dispersal. The strength of compensatory improvements in post-settlement juvenile survival is the most critical factor in determining whether a reserve network can rescue populations from the impacts of severe overharvesting. We use of the model to compare specific alternative proposals for protected area networks along the California coast, as mandated through California’s Marine Life Protection Act, and show that achieving the goals of the Act depends primarily on the fisheries management regulations outside of protected areas, and that the size and configuration of MPAs has a little impact.

Ward, EJ, R Hilborn, RG Towell, L Gerber. 2007. A state–space mixture approach for estimating catastrophic events in time series data. Can. J. Fish. Aquat. Sci. 64:899-910.

Catastrophic events are considered a major contributor to extinction threats, yet rarely included in population viability analysis. We extend the basic state-space population dynamics model to include a mixture distribution for the process error component of the model. The mixture distribution consists of a “normal” component describing regular variability, and a “catastrophic” component. The catastrophic component represents rare events that negatively affect the population. Direct estimation of parameters is rarely possible using a single time series, however estimation is possible when multiple surveys are available, or time series are combined in a meta-analysis. We apply the catastrophic state-space model to simulated time series of abundance from simple non-linear population dynamics models. Applications of the model to these simulated time series indicate that population parameters, and observation and process errors are estimated robustly. Both the frequency and magnitude of catastrophes are susceptible to bias, which is a linear function of the true values of the parameters. Our simulations indicate that the power to detect a catastrophe is also a function of the magnitude of catastrophes, and the degree of observation and process error present. A model that contains a mixture of gamma process errors and gamma observation error is more robust to model misspecification than a model that contains lognormal observation errors.

Branch, TA, R Hilborn, AC Haynie, G Fay, L Flynn, J Griffiths, KN Marshall, JK Randall, JM Scheuerell, EJ Ward, M Young. 2006. Fleet dynamics and fishermen behavior: lessons for fisheries managers. Canadian Journal of Fisheries and Aquatic Sciences 63:1647-1668.

We review fleet dynamics and fishermen behavior from an economic and sociological basis in developing fisheries, in mature fisheries near full exploitation, and in senescent fisheries that are overexploited and overcapitalized. In all cases, fishing fleets behave rationally within the imposed regulatory structures. Successful, generalist fishermen who take risks often pioneer developing fisheries. At this stage, regulations and subsidies tend to encourage excessive entry and investments, creating the potential for serial depletion. In mature fisheries, regulations often restrict season length, vessel and gear types, fishing areas, and fleet size, causing or exacerbating the race for fish and excessive investment, and are typically unsuccessful except when combined with dedicated access privileges (e.g., territorial rights, individual quotas). In senescent fisheries, vessel buyback programs must account for the fishing power of individuals and their vessels. Subsidies should be avoided as they prolong the transition towards alternative employment. Fisheries managers need to create individual incentives that align fleet dynamics and fishermen behavior with the intended societal goals. These incentives can be created both through management systems like dedicated access privileges and through market forces.

Branch, TA, K Rutherford, R Hilborn. 2006. Replacing trip limits with individual transferable quotas: implications for discarding. Mar. Pol. 30:281-292.

Flynn, L, AE Punt, R Hilborn. 2006. A hierarchical model for salmon run reconstruction and application to the Bristol Bay sockeye salmon (Oncorhynchus nerka) fishery. Canadian Journal of Fisheries and Aquatic Sciences 63:1564-1577.

Grafton, R.Q., R. Arnason, T. Bjørndal, D. Campbell, H.F. Campbell, C.W. Clark, R. Connor, D.P. Dupont, R. Hannesson, R. Hilborn, J.E. Kirkley, T. Kompas, D.E. Lane, G.R. Munro, S. Pascoe, D. Squires, S.I. Steinshamn, B.R. Turris, and Q. Weninger. 2006. Incentive-based approaches to sustainable fisheries. Canadian Journal of Fisheries and Aquatic Sciences 63:699-710.

The failures of traditional target-species management have led many to propose an ecosystem approach to fisheries to promote sustainability. The ecosystem approach is necessary, especially to account for fishery-ecosystem interactions, but does little to directly address two important factors of unsustainability — inappropriate incentives for fishers and the ineffective governance that exists in ‘command and control’ fisheries. We contend that much greater emphasis must be placed on fisher motivation when managing fisheries. Using evidence from more than a dozen ‘natural experiments’ in fisheries management, we argue that incentive-based approaches engendered by community, individual harvest or territorial rights, and coupled with public research, monitoring and effective oversight, promote sustainable fisheries.

Keywords: incentives, sustainability, rights, fisheries management

Hilborn, R. 2006. Defining success in fisheries and conflicts in objectives. Mar. Pol. doi:10.1016/j.marpol.2006.05.014/

Hilborn, R. 2006. Faith-based fisheries. Fisheries 31:554-555.

Hilborn, R. 2006. Fisheries success and failure: the case of the Bristol Bay salmon fishery. Bull. Mar. Sci. 78:487-498.

Hilborn, R, P Arcese, P, M Borner, J Hando, G Hopcraft, M Loibooki, S Mduma, ARE Sinclair. 2006. Effective enforcement in a conservation area. Science. 314:1266-1266.

Hilborn, R, J Annala, DS Holland. 2006. The cost of overfishing and management strategies for new fisheries on slow-growing fish: orange roughy (Hoplostethus atlanticus) in New Zealand. Canadian Journal of Fisheries and Aquatic Sciences 63:2149-2153.

Hilborn, R., F. Micheli, and G.A. De Leo. 2006. Integrating marine protected areas with catch regulation. Canadian Journal of Fisheries and Aquatic Sciences 63:642-649.

Previous models of Marine Protected Areas (MPAs) have generally assumed there were no existing regulations on catch, and have frequently shown that MPAs, by themselves, can be used to maintain both sustainable fish stocks and sustainable harvests. We explore the impact of implementing a MPA in a spatially structured model of a single species fish stock that is regulated by total allowable catch (TAC). We find that when a stock is managed at maximum sustainable yield, or is overfished, implementation of a MPA will require a reduction in TAC to avoid increased fishing pressure on the stock outside the MPA. In both cases catches will be lower as a result of overlaying an MPA on existing fisheries management. Only when the stock is so overfished that it is headed towards extinction does a MPA not lead to lower catches. In a TAC regulated fishery, even if the stock is overfished, MPA implementation may not improve overall stock abundance or increase harvest, unless catch is simultaneously reduced in the areas outside the MPA. Models that consider differential adult and larval dispersal need to be explored to see if these results are found with the more complex biology of a two stage model.

Hobbs, N.T., R. Hilborn. Alternatives to statistical hypothesis testing in ecology: A guide to self teaching. Ecological Applications 16:5-19.

Statistical methods emphasizing formal hypothesis testing have dominated the analyses used by ecologists to gain insight from data. Here, we review alternatives to hypothesis testing including techniques for parameter estimation and model selection using likelihood and Bayesian techniques. These methods emphasize evaluation of weight of evidence for multiple hypotheses, multimodel inference, and use of prior information in analysis. We provide a tutorial for maximum likelihood estimation of model parameters and model selection using information theoretics, including a brief treatment of procedures for model comparison, model averaging, and use of data from multiple sources. We discuss the advantages of likelihood estimation, Bayesian analysis, and meta-analysis as ways to accumulate understanding across multiple studies. These statistical methods hold promise for new insight in ecology by encouraging thoughtful model building as part of inquiry, providing a unified framework for the empirical analysis of theoretical models, and by facilitating the formal accumulation of evidence bearing on fundamental questions.

Hodgson, S., Quinn, T.P. Hilborn, R., Francis, R.C. and D.E. Rogers. 2006. Marine and freshwater climatic factors affecting interannual variation in the timing of return migration to fresh water of sockeye salmon (Oncorhynchus nerka). Fisheries Oceanography 14:1-24.

Scheuerell, MD, R Hilborn, MH Ruckelshaus, KK Bartz, KM Lagueux, AD Haas, K Rawson. 2006. The Shiraz model: a tool for incorporating anthropogenic effects and fish–habitat relationships in conservation planning. Canadian Journal of Fisheries and Aquatic Sciences 63:1596-1607.

Current efforts to conserve Pacific salmon (Oncorhynchus spp.) rely on a variety of information sources, including empirical observations, expert opinion, and models. Here we outline a framework for incorporating detailed information on density-dependent population growth, habitat attributes, hatchery operations, and harvest management into conservation planning in a time-varying, spatially explicit manner. We rely on a multistage Beverton ?Holt model to describe the production of salmon from one life stage to the next. We use information from the literature to construct relationships between the physical environment and the necessary productivity and capacity parameters for the model. As an example of how policy makers can use the model in recovery planning, we applied the model to a threatened population of Chinook salmon (Oncorhynchus tshawytscha) in the Snohomish River basin in Puget Sound, Washington, USA. By incorporating additional data on hatchery operations and harvest management for Snohomish River basin stocks, we show how proposed actions to improve physical habitat throughout the basin translate into projected improvements in four important population attributes: abundance, productivity, spatial structure, and life history diversity. We also describe how to adapt the model to a variety of other management applications.

Parma, AM, R Hilborn, JM Orensanz. 2006. The good, the bad and the ugly: learning from experience to achieve sustainable fisheries. Bull. Mar. Sci. 78:411-428.

Branch, TA, R Hilborn, E Bogazzi. 2005. Escaping the tyranny of the grid: a more realistic way of defining fishing opportunities. Canadian Journal of Fisheries and Aquatic Sciences 62:631-642.

de Valpine, P. and R. Hilborn. 2005. State-space likelihoods for nonlinear fisheries time-series. Canadian Journal of Fisheries and Aquatic Sciences 62:1937-1952.

Hyun, S.-Y., Hilborn, R. Anderson J. and Ernst, B. 2005. A statistical model for in-season forecasts of sockeye salmon (Oncorhynchus nerka) returns to the Bristol Bay districts of Alaska. Canadian Journal of Fisheries and Aquatic Sciences 62:1665-1680.

Branch, T. A., Hilborn, R., and Bogazzi, E. 2005. Escaping the tyranny of the grid: a more realistic way of defining fishing opportunities. Canadian Journal of Fisheries and Aquatic Sciences 62:631-642.

Hilborn, R. 2005. Fisheries management. Issues in Science and Technology, 21:10-11.

Hilborn, R., J.M. Orensanz, and A.M. Parma. 2005. Institutions, incentives and the future of fisheries. Philosophical Transactions of the Royal Society of London Series B-Biological Sciences 360:47-57.

Hilborn, R., J.K. Parrish, and K. Litle. 2005. Fishing Rights or Fishing Wrongs? Rev. Fish Biol. Fish. 15:191-198.

Packer, C., R. Hilborn, A. Mosser, B. Kissui, M. Borner, G. Hopcraft, J. Wilmshurst, S. Mduma, and A.R.E. Sinclair. 2005. Ecological Change, Group Territoriality, and Population Dynamics in Serengeti Lions. Science 307:390-393.

Sharma, R., A. Cooper, and R. Hilborn. 2005. A quantitative framework for the analysis of habitat and hatchery practices on Pacific salmon. Ecological Modelling. 183:231-250.

Walters, C.J., and R. Hilborn. 2005. Exploratory assessment of historical recruitment patterns using relative abundance and catch data. Canadian Journal of Fisheries and Aquatic Sciences 62:1985-1990.

Valero, J., C. Hand, J.M. Orensanz, A. M. Parma, D.Armstrong, R. Hilborn. 2004. Geoduck (Panopea abrupta) recruitment trends in the Pacific northwest: long-term changes in relation to climate. California Cooperative Oceanic Fisheries Investigations Report 45:80-86.

Boatright, C., T. Quinn, and R. Hilborn. 2004. Timing of adult migration and stock structure for sockeye salmon in Bear Lake, Alaska. Transactions of the American Fisheries Society 133:911-921.

Orensanz, J. M., C. M. Hand, A. M. Parma, J. Valero, and R. Hilborn. 2004. Precaution in the harvest of Methuselah's clams—the difficulty of getting timely feedback from slowpaced dynamics. Canadian Journal of Fisheries and Aquatic Sciences 61:1355-1372.

Hilborn, R., Punt, A.E., Orensanz, J. 2004. Beyond band-aids in fisheries management: fixing world fisheries. Bulletin of Marine Science 74(3):493-507.

Hilborn, R., K. Stokes, J.J. Maguire, A.D.M. Smith, L.W. Botsford, M. Mangel, J. Orensanz, A. Parma, J. Rice, J. Bell, K.L. Cochrane, S. Garcia, S.J. Hall, G.P. Kirkwood, K. Sainsbury, G. Stefansson, C.J. Walters. 2004. When can marine reserves improve fisheries management? Ocean and Coastal Management 47/3-4:197-205.

Hilborn, R. 2004. Ecosystem-based fisheries management: the carrot or the stick? Marine Ecology-Progress Series 274:275-278.

Flynn, L., and R. Hilborn. 2004. Test fishery indices for sockeye salmon (Oncorhynchus nerka) as affected by age composition and environmental variables. Canadian Journal of Fisheries and Aquatic Sciences 61:80- 92

Gende, S.M., Quinn, T.P., Hilborn, R., Hendry A.P. and Dickerson, B. 2004. Brown bears selectively kill salmon with higher energy content but only in habitats that facilitate choice. Oikos 104:518-528.

Norse, E. A., C. B. Grimes, S. Ralston, R. Hilborn, J. C. Castilla, S. R. Palumbi, D. Fraser, and P. Kareiva. 2003. Marine reserves: the best option for our oceans? Frontiers in Ecology and Environment 1:495- 502.

Hilborn, R., T.A. Branch. B. Ernst, A Magnusson, C.V. Minte-Vera, M.D. Scheuerell, and J.L. Valero. 2003. State of the world's fisheries. Annual Review of Environment and Resources 28:359-399.

Magnusson, A. and R. Hilborn. 2003. Estuarine influence on survival rates of coho (Oncorhynchus kisutch) and chinook salmon (Oncorhynchus tshawytscha) released from hatcheries on the U.S. Pacific coast. Estuaries 26:1094-1103.

Butterworth, D.S., J.N. Ianelli, R. Hilborn. 2003. A statistical model for stock assessment of southern bluefin tuna with temporal changes in selectivity. South African Journal of Marine Science 25:331-361.

Cooper, Andrew B., Hilborn, Ray, Unsworth, James W. 2003. An approach for population assessment in the absence of abundance indices. Ecological Applications 13(3):814-828.

Barrowman, Nicholas J., Myers, Ransom A., Hilborn, Ray, Kehler, Daniel G., Field, Chris A. 2003. The variability among populations of coho salmon in the maximum reproductive rate and depensation. Ecological Applications 13(3):784-793.

Flynn, L., R. Hilborn and A.E. Punt. 2003. Identifying the spatial distribution of stocks of migrating adult sockeye salmon using age composition data. Alaska Fishery Research Bulletin 10:50-60.

Stewart, I.J. R. Hilborn and T.P. Quinn. 2003. Coherence of observed adult sockeye salmon abundance within and among spawning habitats in the Kvichak River watershed. Alaska Fishery Research Bulletin 10:28-41.

Hilborn, R. 2003. The state of the art in stock assessment: where we are and where we are going. Scientia Marina 67(supplement 1):15-20.

Breen, P.A., R. Hilborn, M. Maunder, S. Kim . 2003. Comparing alternative harvest rules to minimise the effects of squid fishery bycatch on Hooker's sea lions (Phocarctos hookeri) in New Zealand. Canadian Journal of Fisheries and Aquatic Sciences 60:527:541.

Hilborn, R., T.P. Quinn, D.E. Schindler and D.E. Rogers. 2003. Biocomplexity and fisheries sustainability. Proceedings of the National Academy of Sciences 100:6564-6568.

Cooper, A. B., J. C. Pinheiro, J.W. Unsworth and R. Hilborn. 2002. Predicting hunter success rates from elk and hunter abundance, season structure, and habitat. Wildlife Society Bulletin 30:1068-1077.

Hilborn, R. 2002. The Dark Side of Reference Points. Bulletin of Marine Science 70:403-408

Myers, R.A., N.J. Barrowman, R. Hilborn and D.G. Kehler. 2002. Inferring Bayesian priors with limited direct data: applications to risk analysis. North American Journal of Fisheries Management 22:351-364.

Hilborn, R. 2002. Marine reserves and fisheries management. Science 295:1233-1234.

Hilborn, R., A. Parma and M. Maunder. 2002. Exploitation rate reference points for west coast rockfish: are they robust and are there better alternatives. North American Journal of Fisheries Management 22:365- 375.

Schindler, D., T.E. Essington, J.F. Kitchell, C. Boggs and R. Hilborn. 2002. Sharks and tunas: fisheries impacts on predators with contrasting life histories. Ecological Applications 12:735-748.

Essington, T.E., J.F. Kitchell, C. Boggs, D.E. Schindler, R.J. Olson and R. Hilborn. 2002. Alternative fisheries and the predation rate of yellowfin tuna in the eastern Pacific Ocean. Ecological Applications 12:724-734.

Rose, K.A., J.H. Cowan Jr., K.O. Winemiller, R.A. Myers and R. Hilborn. 2001. Compensatory density dependence in fish populations: importance, controversy, understanding and prognosis. Fish and Fisheries 2:293-327.

Sharma, R. and R. Hilborn. 2001. Empirical relationships between watershed characteristics and coho salmon (Oncorhynchus kisutch) smolt abundance in 14 western Washington streams. Canadian Journal of Fisheries and Aquatic Sciences 58:1453-1463.

Gerber, L.R. and R. Hilborn. 2001. Catastrophic events and recovery from low densities in populations of otariids: implication for risk of extinction. Mammal Review 11:131-150.

Hilborn, R. and D. Eggers. 2001. A review of the hatchery programs for pink salmon in Prince William Sound and Kodiak Island, Alaska: response to comment. Transactions of the American Fisheries Society 130:720-724.

Liermann, M. and R. Hilborn. 2001. Depensation, evidence, models and implications. Fish and Fisheries 2:33-58.

Hilborn, R. 2001. Calculation of biomass trend, exploitation rate and surplus production from survey and catch data. Canadian Journal of Fisheries and Aquatic Sciences 58:579-584.

Hilborn, R., J.J. Maguire, A. M. Parma, and A. A. Rosenberg. 2001. The precautionary approach and risk management: can they increase the probability of success in fisheries management. Canadian Journal of Fisheries and Aquatic Sciences 58:99-107

Hilborn, R. and D. Eggers. 2000. A review of the hatchery programs for pink salmon in Prince William Sound and Kodiak Island, Alaska. Transactions of the American Fisheries Society 129:333-350.

Hamon, T.R., C.J. Foote, R. Hilborn and D.E. Rogers. 2000. Selection on morphology of spawning wild salmon by a gill-net fishery. Transactions of the American Fisheries Society 129:1300-1315.

Maunder, M., Starr, P.J. and R. Hilborn. 2000. A Bayesian analysis to estimate loss in squid catch due to the implementation of a sea lion population management plan. Marine Mammal Science 16: 413-426.

Mduma, S. Sinclair, A.R.E. and R. Hilborn. 1999. Food regulates the Serengeti wildebeest: a forty-year record. Journal of Animal Ecology 68:1101-1122.

Hilborn, R., B.G. Bue, and S. Sharr. 1999. Estimating spawning escapements from periodic counts: a comparison of methods. Canadian Journal of Fisheries and Aquatic Sciences 56:888-896.

Hilborn, R. 1999. Confessions of a reformed hatchery basher. Fisheries 24:30-31.

Hilborn, R. and M. Liermann. 1998. Standing on the shoulders of giants: learning from experience. Reviews in Fish Biology and Fisheries 8:273-283

Coronado, C. and R. Hilborn. 1998. Spatial and temporal factors affecting survival in coho salmon (Oncorhynchus kisutch) in the Pacific Northwest. Canadian Journal of Fisheries and Aquatic Sciences 55:2067-2077.

Hilborn, R. 1998. The economic performance of marine stock enhancement projects. Bulletin of Marine Science 62:661-674

Coronado, C. and R. Hilborn. 1998. Spatial and temporal factors affecting survival in coho and fall chinook salmon in the Pacific northwest. Bulletin of Marine Science 62:409-425.

Orensanz, J.M., J. Armstrong, D. Armstrong and R. Hilborn. 1998. Crustacean resources are vulnerable to serial depletion—the multifaceted declines of crab and shrimp fisheries in the Greater Gulf of Alaska. Reviews in Fish Biology and Fisheries 8:117-176

Starr, P., J.H. Annala and R. Hilborn. 1998. Contested stock assessment: two case studies. Canadian Journal of Fisheries and Aquatic Sciences 55:529-537.

Prince J. and R. Hilborn. 1998. Concentration profiles and invertebrate fisheries management.. Canadian Special Publication of Fisheries and Aquatic Sciences 125:187-196.

Hilborn, R. 1997. Statistical hypothesis testing and decision theory in fisheries science. Fisheries 22(10):19-20

Pascual, M.A., P. Kareiva and R. Hilborn. 1997. The influence of model structure on conclusions about the viability and harvesting of Serengeti wildebeest. Conservation Biology 11:966-976.

Starr, P.J., Breen, P.A., Hilborn, R. and T.H. Kendrick. 1997. Evaluation of a management decision rule for a New Zealand rock lobster substock. Mar. Freshwater Res. 48:1093-1101.

Hilborn, R. 1997. Lobster stock assessment: report from a workshop; II. Mar. Freshwater Res. 48:945-947

Liermann, M. and R. Hilborn. 1997. Depensation in fish stocks: a hierarchic Bayesian meta-analysis. Canadian Journal of Fisheries and Aquatic Sciences 54:1976-1984.

Fogarty, M.J., Hilborn, R. and D. Gunderson. 1997. Chaos and parametric management. Marine Policy 21:187-194.

Punt, A.E. and R. Hilborn. 1997. Fisheries stock assessment and decision analysis: the Bayesian approach. Reviews in Fish Biology and Fisheries 7:35-63.

Hilborn, R. 1997. Recruitment paradigms for fish stocks. Canadian Journal of Fisheries and Aquatic Sciences 54:984-985

Hilborn, R. 1996. Risk analysis in fisheries and natural resource management. Human Ecology and Risk Assessment 2:655-659.

Hilborn, R. 1996. Do principles for conservation help managers? Ecological Applications 6:364-365.

Hilborn, R. and D. Gunderson. 1996. Chaos and paradigms for fisheries management. Marine Policy 20:87-89.

Hilborn, R., C.J. Walters and D. Ludwig. 1995. Sustainable exploitation of renewable resources. Annual Review of Ecology and Systematics 26:45- 67.

Pascual, M. A., and R. Hilborn. 1995. Conservation of harvested populations in fluctuating environments: the case of the Serengeti wildebeest. Journal of Applied Ecology 32:468-480.

McAllister, M.M, E.K. Pikitch, A.E. Punt and R. Hilborn. 1994. A Bayesian approach to stock assessment and harvest decisions using the sampling/importance resampling algorithm. Canadian Journal of Fisheries and Aquatic Sciences 51:2673-2687.

Anganuzzi, A., R. Hilborn and J. R. Skalski. 1994. Estimation of size selectivity and movement rates from mark-recapture data. Canadian Journal of Fisheries and Aquatic Sciences 51:734-742

Punt, A.E. and R. Hilborn. 1994. A comparison of fishery models with and without cannibalism with implications for the management of the Cape hake resource off southern Africa. ICES Journal of Marine Science 51:19-29

Winton, J. and R. Hilborn. 1994. Lessons from supplementation of chinook salmon in British Columbia. North American Journal of Fisheries Management 14:1-13

Hilborn, R., E. K. Pikitch, and M. K. McAllister. 1994. A Bayesian estimation and decision analysis for an age-structured model using biomass survey data. Fisheries Research 19:17-30

Schnute, J. T. and R. Hilborn. 1993. Analysis of contradictory data sources in fish stock assessment. Canadian Journal of Fisheries and Aquatic Sciences 50:1916-1923

Polacheck, T., R. Hilborn and A. E. Punt. 1993. Fitting surplus production models: comparing methods and measuring uncertainty. Canadian Journal of Fisheries and Aquatic Sciences 50:2597-2607.

Hilborn, R. and J. Winton. 1993. Learning to enhance salmon production: lessons from the salmonid enhancement program. Canadian Journal of Fisheries and Aquatic Sciences 50:2043-2056

Hilborn, R. and D. Ludwig. 1993. The limits of applied ecological research. Ecological Applications 3:550-552

Ludwig, D., R. Hilborn, and C. Walters. 1993. Uncertainty, resource exploitation, and conservation: lessons from history. Science 260:17/36.

Hilborn, R., E. K. Pikitch, M. K. McAllister, and A. E. Punt. 1993. Use of Risk Analysis to Assess Fishery Management Strategies—a Case-Study Using Orange Roughy (Hoplostethus atlanticus) on the Chatham Rise, New-Zealand—Comment. Canadian Journal of Fisheries and Aquatic Sciences 50:1122-1125.

Hilborn, R., E.K. Pikitch, and R.C. Francis. 1993. Current trends in including risk and uncertainty in stock assessment and harvest decisions. Canadian Journal of Fisheries & Aquatic Sciences 50:874-880.

Hilborn, R. 1992. Current and future trends in fisheries stock assessment and management. South African Journal of Marine Science 12:975-988.

Hilborn, R. 1992. Can fisheries agencies learn from experience? Fisheries 17:6-14.

Hilborn, R. 1992. Hatcheries and the future of salmon in the northwest. Fisheries 17:5-8.

Hilborn, R. 1992. Institutional learning and spawning channels for sockeye salmon (Oncorhynchus nerka). Canadian Journal of Fisheries and Aquatic Science 49:1126-1136.

Hilborn, R. and R. Kennedy. 1992. Spatial pattern in catch rates: a test of economic theory. Bulletin of Mathematical Biology 54:263-273.

Hilborn, R., and C. J. Krebs. 1992. Bias in the Minimum Number Alive Estimator - a Reply. Canadian Journal of Zoology. 70:632-632.

Hilborn, R. 1991. Modeling the stability of fish schools: exchange of individual fish between schools of skipjack tuna (Katsuwonus pelamis). Canadian Journal of Fisheries and Aquatic Sciences 48:1081-1091.

Hilborn, R. 1990. Determination of fish movement patterns from tag recoveries using maximum likelihood estimators. Canadian Journal of Fisheries and Aquatic Sciences 47:635-643.

Hilborn, R. 1989. Yield estimation for spatially connected populations: an example of surface and longline fisheries for yellowfin tuna. North American Journal of Fisheries Management 9:402-410.

Hilborn, R. 1989. Models of tag dynamics with exchange between available and unavailable populations. Canadian Journal of Fisheries and Aquatic Sciences 46:1356-1366.

Hilborn, R. 1989. International Management of Tuna. Marine Policy, 13:166- 166.

Hilborn, R. and P. Medley. 1989. Tuna purse seine fishing with fish aggregating devices: models of tuna FAD interactions. Canadian Journal of Fisheries and Aquatic Sciences 46:28-32.

Hilborn, R. and J. Sibert. 1988. Is international management of tuna necessary? Marine Policy. January 1988, pp. 31-39.

Hilborn, R. and J. Sibert. 1988. Adaptive management of developing fisheries. Marine Policy. April 1988, pp. 112-121.

Hilborn, R. 1988. Determination of tag return from recaptured fish by sequential examination for tags. Transactions of the American Fisheries Society 117:510-514.

Starr, P.J. and R. Hilborn. 1988. Reconstruction of harvest rates and stock contribution in gauntlet salmon fisheries. Canadian Journal of Fisheries and Aquatic Sciences 45:2216-2229.

Hall, D.L, R. Hilborn, M. Stocker and C.J. Walters. 1988. Alternative harvest strategies for Pacific Herring (Clupea harengus pallasi). Canadian Journal of Fisheries and Aquatic Sciences 45:888-897.

Fried, S.M. and R. Hilborn. 1988. A Bayesian approach to inseason run size estimation for Bristol Bay, Alaska, Sockeye Salmon (Oncorhynchus nerka). Canadian Journal of Fisheries and Aquatic Sciences 45:850-855.

Hilborn, R. and W. Luedke. 1987. Rationalizing the irrational: a case study in user group participation in Pacific salmon management. Canadian Journal of Fisheries and Aquatic Sciences 44:1796-1805.

Hilborn, R. 1987. Living with uncertainty in resource management. North American Journal of Fisheries Management 7:1-5.

Shardlow, T., R. Hilborn, and D. Lightly. 1987. Components analysis of instream escapement methods for Pacific Salmon. Canadian Journal of Fisheries and Aquatic Sciences 44:1031-1037.

Hilborn, R. and C.J. Walters. 1987. A general model for simulation of stock and fleet dynamics in spatially heterogeneous fisheries. Canadian Journal of Fisheries and Aquatic Sciences 44:1366-1370.

Hilborn, R. and C.J. Walters. 1987. Microcomputer simulation for training and teaching. Environmental Software 1:156-163.

Hilborn, R. 1986. A comparison of alternative harvest tactics for invertebrate fisheries. Pp. 313-317 in G.S. Jamieson and N. Bourne (eds.), Canadian Special Publication in Fisheries and Aquatic Sciences 92.

Hilborn, R. 1986. Some Remarks on Determinants of Catching Power in the British-Columbia Salmon Purse Seine Fleet by Hilborn and Ledbetter—Reply. Canadian Journal of Fisheries and Aquatic Sciences, 43:1086- 1088.

Moussalli, E. and R. Hilborn. 1986. Optimal stock size and harvest rate in multistage life history models. Canadian Journal of Fisheries and Aquatic Sciences 43:135-141.

Renyard, T.S. and R. Hilborn. 1986. Sport angler preferences for alternative regulatory methods. Canadian Journal of Fisheries and Aquatic Sciences 43:240-242.

Lawson, T.A. and R. Hilborn. 1985. Equilibrium yields and yield isopleths from a general age-structured model of harvested populations. Canadian Journal of Fisheries and Aquatic Sciences 42:1766-1771.

Hilborn, R. 1985. Simplified calculation of optimum spawning stock size from Rickers' stock recruitment curve. Canadian Journal of Fisheries and Aquatic Sciences 42:1834-1835.

Hilborn, R. 1985. Fleet dynamics and individual variation: why some people catch more fish than others. Canadian Journal of Fisheries and Aquatic Sciences 42:2-13.

Hilborn, R. 1985. Apparent stock recruitment relationships in mixed stock fisheries. Canadian Journal of Fisheries and Aquatic Sciences 42:718-723.

Shardlow, T. and R. Hilborn. 1985. Density dependent catchability coefficients. Transactions of the American Fisheries Society 114:436-438.

Hilborn, R. and M. Ledbetter. 1985. Determinants of catching power in the B.C. salmon purse seine fleet. Canadian Journal of Fisheries and Aquatic Sciences 42:51-56.

Hilborn, R., C.J. Walters, R.M. Peterman and M.J. Staley. 1984. Models and fisheries: A case study in implementation. North American Journal of Fisheries Management 4:9-15.

Walker, K.D., R.B. Rettig, and R. Hilborn. 1983. Analysis of multiple objectives in Oregon coho salmon policy. Canadian Journal of Fisheries and Aquatic Sciences 40:580-587.

Argue, A.W., R. Hilborn, R.M. Peterman, M.J. Staley, and C.J. Walters. 1983. The Strait of Georgia Chinook and Coho fishery. Canadian Journal of Fisheries and Aquatic Sciences Bulletin 211.

Ludwig, D. and R. Hilborn. 1983. Adaptive probing strategies for age structured fish stocks. Canadian Journal of Fisheries and Aquatic Sciences 40:559-569.

Stocker, M. and R. Hilborn. 1982. Comment on "Short-term forecasting in marine fish stocks." Canadian Journal of Fisheries and Aquatic Sciences 39:1071-1072.

Hilborn, R. and S. Stearns. 1982. On inference in ecology and evolutionary biology: the problem of multiple causes. Acta Biotheoretica 32:145-164.

Hilborn, R. and C.J. Walters. 1981. Pitfalls of environmental baseline and process studies. Environmental Impact Assessment Review 2: 265-278.

Stocker, M. and R. Hilborn. 1981. Short term forecasting in marine fish stocks. Canadian Journal of Fisheries and Aquatic Sciences 38:1247-1254.

Yom-Tov, Y. and R. Hilborn. 1981. Energetic constraints on clutch size and time of breeding in temperate zone birds. Oecologia 48:234-243.

Hilborn, R. 1979. Comparison of fisheries control systems that utilize catch and effort data. Journal of the Fisheries Research Board of Canada 36:1477-1489.

Hilborn, R. 1979. Some failures and successes at applying systems analysis to ecological problems. Journal of Applied Systems Analysis 6:25-31.

Hilborn, R. and M. Ledbetter. 1979. Analysis of the British Columbia salmon purse seine fleet: dynamics of movement. Journal of the Fisheries Research Board of Canada 36:384-391.

Hilborn, R. 1979. Some long term dynamics of predator–prey models with diffusion. Ecological Modelling 6:23-30.

Walters, C.J., R. Hilborn, R.M. Peterman and M. Staley. 1978. A model for examining early ocean limitation of Pacific salmon production. Journal of the Fisheries Research Board of Canada 35:1303-1315.

Walters, C.J. and R. Hilborn. 1978. Ecological optimization and adaptive management. Annual Review of Ecology and Systematics 8:157-188.

Hilborn, R. and C.J. Walters. 1977. Differing goals of salmon management on the Skeena River. Journal of the Fisheries Research Board of Canada 34:64-72.

Hilborn, R., C.J. Krebs, and J.A. Redfield. 1976. On the reliability of enumeration for mark and recapture census of voles. Canadian Journal of Zoology 54:1019-1024.

Hilborn, R. and C.J. Krebs. 1976. Fates of disappearing individuals in fluctuating populations of Microtus townsendi. Canadian Journal Zoology 54:1501-1520.

Hilborn, R. 1976. Optimal exploitation of multiple stocks by a common fishery. Journal of the Fisheries Research Board of Canada 33:1-5.

Walters, C.J. and R. Hilborn. 1976. Adaptive control of fishing systems. Journal of the Fisheries Research Board of Canada 33:145-159.

Krebs, C.J. I. Wingate, J. LeDuc, J. Redfield, M. Taitt and R. Hilborn. 1976. Microtus population biology: dispersal in fluctuating populations of Microtus townsendi. Canadian Journal of Zoology 54:79-95.

Hilborn, R. 1975. Similarities in dispersal tendency among siblings in four species of voles (Microtus). Ecology 56:1221-1225.

Hilborn, R. 1975. The effect of spatial heterogeneity on the persistence of predator-prey interactions. Theoretical Population Biology 8:346-355.

Walters, C.J., Bunnell, F., Hilborn, R., and R.M. Peterman. 1975. Computer simulation of barren ground caribou dynamics. Ecological Modelling 1:303-315.

Himamowa, Bubu. 1975. The Obergurgl model: a microcosm of economic growth in relation to limited ecological resources. Nature and Resources 2:10-21. (Himamowa is a pseudonym for myself and six other authors)

Hilborn, R. 1974. Inheritance of skeletal polymorphism in Microtus californicus. Heredity 33:81-83.

Walters, C.J. R. Hilborn, E. Oguss, R. Peterman and J. Stander. 1974. Development of a simulation model of Mallard duck populations. Canadian Wildlife Service Occasional Paper No. 20:1-34.

Hilborn, R. 1973. A control system for FORTRAN simulation programming. Simulation. 20:172-175.