Age-dependent survival analyzed with Bayesian models of mark-recapture data

We describe a Bayesian random effects model of mark-recapture data that accounts for age-dependence in survival and individual heterogeneity in capture probabilities and survival. The model is applied to data on the Glanville fritillary butterfly (Melitaea cinxia) collected from a population enclosed in a large cage in the field. The cage population consisted of a mixture of butterflies originating from newly established and old populations in a large metapopulation in the Aland Islands in Finland. The explanatory variables in the model included the effects of temperature, sex, and population type (new vs. old) on capture probabilities, and the effects of age, sex, population type, and day vs. night on survival. We found that mortality rate increased with age, that mortality rate was much higher during the day than during the night, and that the life span of females originating from newly established populations was shorter than the life span of females from old populations. Capture probability decreased with increasing temperature and decreased with increasing mobility of individuals.     

Multiple data sources improve DNA-based mark-recapture population estimates of grizzly bears.

A fundamental challenge to estimating population size with mark-recapture methods is heterogeneous capture probabilities and subsequent bias of population estimates. Confronting this problem usually requires substantial sampling effort that can be difficult to achieve for some species, such as carnivores. We developed a methodology that uses two data sources to deal with heterogeneity and applied this to DNA mark-recapture data from grizzly bears (Ursus arctos). We improved population estimates by incorporating additional DNA "captures" of grizzly bears obtained by collecting hair from unbaited bear rub trees concurrently with baited, grid-based, hair snag sampling. We consider a Lincoln-Petersen estimator with hair snag captures as the initial session and rub tree captures as the recapture session and develop an estimator in program MARK that treats hair snag and rub tree samples as successive sessions. Using empirical data from a large-scale project in the greater Glacier National Park, Montana, USA, area and simulation modeling we evaluate these methods and compare the results to hair-snag-only estimates. Empirical results indicate that, compared with hair-snag-only data, the joint hair-snag-rub-tree methods produce similar but more precise estimates if capture and recapture rates are reasonably high for both methods. Simulation results suggest that estimators are potentially affected by correlation of capture probabilities between sample types in the presence of heterogeneity. Overall, closed population Huggins-Pledger estimators showed the highest precision and were most robust to sparse data, heterogeneity, and capture probability correlation among sampling types. Results also indicate that these estimators can be used when a segment of the population has zero capture probability for one of the methods. We propose that this general methodology may be useful for other species in which mark-recapture data are available from multiple sources.     

Estimating species-specific survival and movement when species identification is uncertain

Incorporating uncertainty in the investigation of ecological studies has been the topic of an increasing body of research. In particular, mark-recapture methodology has shown that incorporating uncertainty in the probability of detecting individuals in populations enables accurate estimation of population-level processes such as survival, reproduction, and dispersal. Recent advances in mark-recapture methodology have included estimating population-level processes for biologically important groups despite the misassignment of individuals to those groups. Examples include estimating rates of apparent survival despite less than perfect accuracy when identifying individuals to gender or breeding state. Here we introduce a method for estimating apparent survival and dispersal in species that co-occur but that are difficult to distinguish. We use data from co-occurring populations of meadow voles (Microtus pennsylvanicus) and montane voles (M. montanus) in addition to simulated data to show that ignoring species uncertainty can lead to biased estimates of population processes. The incorporation of species uncertainty in mark-recapture studies should aid future research investigating ecological concepts such as interspecific competition, niche differentiation, and spatial population dynamics in sibling species.     

A multi-event model to study stage-dependent dispersal in radio-collared hares: when hunting promotes costly transience

Behavioral ecologists have often assumed that dispersal is costly mainly because of unfamiliarity with traversed habitats during dispersal and energy costs of the movement per se; thus, dispersers that have successfully settled should experience survival rates comparable to those of philopatric individuals. In this paper, we tested that hypothesis using 152 radio-collared European hares in a harvested population. We developed a multi-event capture recapture model, combining telemetry data and recoveries and separately modeling the foray probability, the settlement probability, and the permanent dispersal probability. The parameterization introduced here raises the possibility of separately testing effects on survival and dispersal probabilities at each stage of dispersal (departure, transience, and settlement). In accordance with our expectations, we reveal that dispersers incur higher mortality risks during transience and the early settlement period than philopatric individuals or settled dispersers. We also found that dispersers suffer from higher risks of being shot. Those results illustrate that unfamiliarity with the habitat during transience makes dispersal costly and that settled dispersers may enjoy survival rates comparable to those of philopatric individuals. Surprisingly, we also found that individuals have a higher probability of foraying during the hunting season. We suggest that hunting and related disturbances increase dispersal costs both by increasing mortality risk during transience and (perhaps) by increasing movement rates. We emphasize the need to take human pressures into account as factors that may drive the demographics of movements in populations.     

Photographic mark-recapture analysis of local dynamics within an open population of dolphins

Identifying demographic changes is important for understanding population dynamics. However, this requires long-term studies of definable populations of distinct individuals, which can be particularly challenging when studying mobile cetaceans in the marine environment. We collected photo-identification data from 19 years (1992-2010) to assess the dynamics of a population of bottlenose dolphins (Tursiops truncatus) restricted to the shallow (<7 m) waters of Little Bahama Bank, northern Bahamas. This population was known to range beyond our study area, so we adopted a Bayesian mixture modeling approach to mark-recapture to identify clusters of individuals that used the area to different extents, and we specifically estimated trends in survival, recruitment, and abundance of a "resident" population with high probabilities of identification. There was a high probability (p= 0.97) of a long-term decrease in the size of this resident population from a maximum of 47 dolphins (95% highest posterior density intervals, HPDI = 29-61) in 1996 to a minimum of just 24 dolphins (95% HPDI = 14-37) in 2009, a decline of 49% (95% HPDI = approximately 5% to approximately 75%). This was driven by low per capita recruitment (average approximately 0.02) that could not compensate for relatively low apparent survival rates (average approximately 0.94). Notably, there was a significant increase in apparent mortality (approximately 5 apparent mortalities vs. approximately 2 on average) in 1999 when two intense hurricanes passed over the study area, with a high probability (p = 0.83) of a drop below the average survival probability (approximately 0.91 in 1999; approximately 0.94, on average). As such, our mark-recapture approach enabled us to make useful inference about local dynamics within an open population of bottlenose dolphins; this should be applicable to other studies challenged by sampling highly mobile individuals with heterogeneous space use.     

Science or Slaughter: Need for Lethal Sampling of Sharks

Abstract: General consensus among scientists, commercial interests, and the public regarding the status of shark populations is leading to an increasing need for the scientific community to provide information to help guide effective management and conservation actions. Experience from other marine vertebrate taxa suggests that public, political, and media pressures will play an increasingly important part in setting research, management, and conservation priorities. We examined the potential implications of nonscientific influences on shark research. In particular, we considered whether lethal research sampling of sharks is justified. Although lethal sampling comes at a cost to a population, especially for threatened species, the conservation benefits from well‐designed studies provide essential data that cannot be collected currently in any other way. Methods that enable nonlethal collection of life‐history data on sharks are being developed (e.g., use of blood samples to detect maturity), but in the near future they will not provide widespread or significant benefits. Development of these techniques needs to continue, as does the way in which scientists coordinate their use of material collected during lethal sampling. For almost half of the known shark species there are insufficient data to determine their population status; thus, there is an ongoing need for further collection of scientific data to ensure all shark populations have a future. Shark populations will benefit most when decisions about the use of lethal sampling are made on the basis of scientific evidence that is free from individual, political, public, and media pressures.     

Contrasting diel patterns in vertical movement and locomotor activity of whale sharks at Ningaloo Reef

Activity patterns of animals often relate to environmental variables such as food availability and predation pressure. Technological advances are providing us with new tools to monitor and better understand these activity patterns. We used animal-attached data loggers recording acceleration and depth to compare activity patterns and vertical habitat use of whale sharks (Rhincodon typus) at Ningaloo Reef, Western Australia. Whale sharks showed a moderate reverse diel vertical migration but exhibited a clear crepuscular pattern in locomotory activity. Peak activity occurred at sunset, whereas vertical movement peaked prior to this. Typical ram surface filter feeding could be identified and occurred primarily during sunset and the first hours of night. At such times, direct observations indicated whale sharks were feeding on tropical krill swarms. Kinematic analysis of postural data and data from vertical movement suggests that whale sharks at Ningaloo spend ~8 min per day actively ram surface filter feeding. Considering the high biomass present in krill schools, it is estimated that whale sharks at Ningaloo have a similar energy intake as those at other aggregation sites. Diel patterns in activity and diving behaviour suggest that whale sharks have tuned their diving behaviour in anticipation of the formation of these high-density patches which appear to only be periodically, but predictably available at sunset. Our results confirm that diel patterns in vertical habitat selection and vertical movements do not necessarily reflect patterns in activity and foraging behaviour. Direct quantification of activity and behaviour is required in gaining accurate representation of diel activity patterns.     

Estimating survival rates of uncatchable animals: the myth of high juvenile mortality in reptiles

Survival rates of juvenile reptiles are critical population parameters but are difficult to obtain through mark-recapture programs because these small, secretive animals are rarely caught. This scarcity has encouraged speculation that survival rates of juveniles are very low, and we test this prediction by estimating juvenile survival rates indirectly. A simple mathematical model calculates the annual juvenile survival rate needed to maintain a stable population size, using published data on adult survival rates, reproductive output, and ages at maturity in 109 reptile populations encompassing 57 species. Counter to prediction, estimated juvenile survival rates were relatively high (on average, only about 13% less than those of conspecific adults) and highly correlated with adult survival rates. Overall, survival rates during both juvenile and adult life were higher in turtles than in snakes, and higher in snakes than in lizards. As predicted from life history theory, rates of juvenile survival were higher in species that produce large offspring, and higher in viviparous squamates than in oviparous species. Our analyses challenge the widely held belief that juvenile reptiles have low rates of annual survival and suggest instead that sampling problems and the elusive biology of juvenile reptiles have misled researchers in this respect.     

Long-term individual identification and site fidelity of white sharks, <Emphasis Type="Italic">Carcharodon carcharias</Emphasis>, off California using dorsal fins

Mark-recapture techniques can be used to estimate white shark (Carcharodon carcharias) population abundance. These frameworks are based on assumptions that marks are conserved and animals are present at the sampling location over the entire duration of the study. Though these assumptions have been validated across short-time scales for white sharks, long-term studies of population trends are dependent on these assumptions being valid across longer periods. We use 22 years of photographic data from aggregation sites in central California to support the use of dorsal fin morphology as long-term individual identifiers. We identified five individuals over 16–22 years, which support the use of dorsal fins as long-time individual identifiers, illustrate strong yearly site fidelity to coastal aggregation sites across extended time periods (decades), and provide the first empirical validation of white shark longevity >22 years. These findings support the use of fin morphology in mark-recapture frameworks for white sharks.     

A generalized Chao estimator with measurement error and external information

We present a generalized Chao (GC) estimator based on a subject-occasion-specific design matrix. We then extend the GC estimator to (i) external information, in the form of non-linear constraints on subpopulation sizes and (ii) measurement error. For the first, we propose a reparameterization of the estimating equations. As a result, the constrained MLE can be found with no additional computational efforts. For the second we generalize SIMEX procedure to multiple measurement methods. In simulation we show that (even incorrect) external information can substantially decrease the MSE. We illustrate with an application to a whale shark (Rhincodon typus) population, where mostly jouvenile males are observed. We use external information on gender ratio of whale sharks to correct for low catchability of females, and our multivariate SIMEX procedure to correct for measurement error in assessment of shark length. The resulting population size estimates are about 60% larger than the unconstrained–uncorrected counterparts.     

Potential effects of environmental contamination on Yuma Myotis demography and population growth.

Unplanned natural and anthropogenic disasters provide unique opportunities for investigating the influence of perturbations on population vital rates and species recovery times. We investigated the potential effects of a major pesticide spill by comparing annual survival rates using mark-recapture techniques on a riparian bat species, Yuma Myotis (Myotis yumanensis). Demography and population dynamics for most bat species remain poorly understood despite advances in mark-recapture estimation and modeling techniques. We compared survival and population growth rates of two roost populations exposed to a large chemical (metam sodium) spill in the upper Sacramento River in Northern California with two roost populations outside the contaminated area from 1992 to 1996. Hypotheses about long-term effects of the spill on female juvenile and adult survival were tested using an information-theoretic approach (AIC). Working hypotheses included effects of age, chemical spill, and time trend on survival. Female adult survival was higher than female juvenile survival across all sites, suggesting stage-specific mortality risks. Model-averaged estimates of female juvenile survival in the contaminated area (0.50-0.74) were lower than in control roosts (0.60-0.78) for each year in the study, suggesting that the spill may have reduced juvenile survival for several years. Female adult survival (0.73-0.89) did not appear to be strongly affected by the spill during the years of the study. There was an increase in survival for both stage-classes across all populations during the study period, which may have been caused by the end of an extended drought in California in the winter of 1993. The spill-affected population was in decline for the first year of the study as indicated by an estimated growth rate (lambda) < 1, but population growth rates increased during the four-year period.     

Predicting occurrence of juvenile shark habitat to improve conservation planning

Fishing and habitat degradation have increased the extinction risk of sharks, and conservation strategies recognize that survival of juveniles is critical for the effective management of shark populations. Despite the rapid expansion of marine protected areas (MPAs) globally, the paucity of shark‐monitoring data on large scales (100s–1000s km) means that the effectiveness of MPAs in halting shark declines remains unclear. Using data collected by baited remote underwater video systems (BRUVS) in northwestern Australia, we developed generalized linear models to elucidate the ecological drivers of habitat suitability for juvenile sharks. We assessed occurrence patterns at the order and species levels. We included all juvenile sharks sampled and the 3 most abundant species sampled separately (grey reef [Carcharhinus amblyrhynchos], sandbar [Carcharhinus plumbeus], and whitetip reef sharks [Triaenodon obesus]). We predicted the occurrence of juvenile sharks across 490,515 km² of coastal waters and quantified the representation of highly suitable habitats within MPAs. Our species‐level models had higher accuracy (? ≥ 0.69) and deviance explained (≥48%) than our order‐level model (? = 0.36 and deviance explained of 10%). Maps of predicted occurrence revealed different species‐specific patterns of highly suitable habitat. These differences likely reflect different physiological or resource requirements between individual species and validate concerns over the utility of conservation targets based on aggregate species groups as opposed to a species‐focused approach. Highly suitable habitats were poorly represented in MPAs with the most restrictions on extractive activities. This spatial mismatch possibly indicates a lack of explicit conservation targets and information on species distribution during the planning process. Non‐extractive BRUVS provided a useful platform for building the suitability models across large scales to assist conservation planning across multiple maritime jurisdictions, and our approach provides a simple for method for testing the effectiveness of MPAs.     

Edge Effects and Isolation: Red-Backed Voles Revisited

Abstract:  We examined demographic responses of California red-backed voles ( Clethrionomys californicus ) to forest fragmentation in southwestern Oregon at sites where this species has previously shown negative responses to fragmentation. Voles were captured in live traps and released. Voles were rarely caught in clearcuts surrounding 11 forest fragments, but relative vole density did not decrease from the forest-fragment interiors to edges. The first result agrees with previous findings at these sites 6 years earlier, but the latter result does not. There was no evidence that vole response to edge changes with fragment age. Two years of intensive mark-recapture efforts at two forest-fragment sites and two unfragmented (control) sites did not show negative effects of fragmentation on vole survival, an important demographic rate. Vole capture probabilities varied greatly across space and time on these four sites, which may explain the differences in vole responses to edge seen between this and the previous study. These results suggest that reliable appraisal of edge effects may be difficult for many species on small fragments because the data necessary to apply population estimators require great efforts to obtain and the use of indices leads to a confounding of detection probabilities with demographic change.     

Protected species use of a coastal marine migratory corridor connecting marine protected areas

The establishment of protected corridors linking the breeding and foraging grounds of many migratory species remains deficient, particularly in the world’s oceans. For example, Australia has recently established a network of Commonwealth Marine Reserves, supplementing existing State reserves, to protect a wide range of resident and migratory marine species; however, the routes used by mobile species to access these sites are often unknown. The flatback marine turtle (Natator depressus) is endemic to the continental shelf of Australia, yet information is not available about how this species uses the marine area. We used a geospatial approach to delineate a coastal corridor from 73 adult female flatback postnesting migratory tracks from four rookeries along the north-west coast of Australia. A core corridor of 1,150 km length and 30,800 km² area was defined, of which 52 % fell within 11 reserves, leaving 48 % (of equivalent size to several Commonwealth Reserves) of the corridor outside of the reserve network. Despite limited data being available for other marine wildlife in this region, humpback whale migratory tracks overlapped with 96 % of the core corridor, while the tracks of three other species overlapped by 5–10 % (blue whales, olive ridley turtles, whale sharks). The overlap in the distribution ranges of at least 20 other marine vertebrates (dugong, cetaceans, marine turtles, sea snakes, crocodiles, sharks) with the corridor also imply potential use. In conclusion, this study provides valuable information towards proposing new locations requiring protection, as well as identifying high-priority network linkages between existing marine protected areas.     

Individual heterogeneity in recapture probability and survival estimates in cheetah

Accurate estimates of demographic parameters are key for understanding and predicting population dynamics and for providing insights for effective wildlife management. Up until recently, no suitable methodology has been available to estimate survival probabilities of species with asynchronous reproduction and a high level of individual variation in capture probabilities. The present work develops a capture-mark-recapture model for cheetahs in the Serengeti National Park, Tanzania, which (a) deals with continuous reproduction, (b) takes into account the high level of individual heterogeneity in capture probabilities and (c) is spatially explicit. Results show that (1) our approach, which is an extensive modification of the Cormack-Jolly-Seber model, provides a lower female adult survival estimate and a higher male adolescent survival estimate than previous approaches to estimate cheetah survival in the area, (2) using sighting location alone is not sufficient to capture the individual variation in resighting probabilities for both sexes, and (3) precision in estimated survival probabilities is generally increased. Species which are individually recognizable, wide-ranging and/or where individuals differ substantially in sightability are particularly appropriate to our modelling approach, and our methodology would thus be appropriate for a wide number of species to provide more accurate estimates of survival.     

Increasing the Accuracy of Productivity and Survival Estimates in Assessing Landbird Population Status

Abstract:  The conservation of species with declining populations requires information on population demography and identification of factors that limit population growth. For landbird species, an understanding of large-scale population declines often requires assessment of local population processes, including the production of offspring, the survival of those offspring, and adult survival. Population growth has been modeled for several species of landbirds to date, and these studies have provided important information on relationships between population status and population-limiting factors. Several recent studies have illuminated field methods and analytical techniques that can aid in increasing the accuracy of productivity and survival estimates for population models. We reviewed these methods and recommend their implementation, including quantification of the season-long productivity of individuals, collection of empirical data on juvenile survival during the postfledging and overwintering periods, and incorporation of adult breeding dispersal into annual adult survival estimates. Such methods will allow for more accurate assessment of population status and provide a better understanding of the factors on which to focus our conservation efforts.     

Habitat,movements and environmental preferences of dusky sharks,Carcharhinus obscurus,in the northern Gulf of Mexico

The dusky shark (Carcharhinus obscurus) is the largest member of the genus Carcharhinus and inhabits coastal and pelagic ecosystems circumglobally in temperate, subtropical and tropical marine waters. In the western North Atlantic Ocean (WNA), dusky sharks are overfished and considered vulnerable by the International Union for the Conservation of Nature. As a result, retention of dusky sharks in commercial and recreational fisheries off the east coast of the United States (US) and in the northern Gulf of Mexico is prohibited. Despite the concerns regarding the status of dusky sharks in the WNA, little is known about their habitat utilization. During the summers of 2008–2009, pop-up satellite archival tags were attached to ten dusky sharks (one male, nine females) at a location where they have been observed to aggregate in the north central Gulf of Mexico southwest of the Mississippi River Delta to examine their movement patterns and habitat utilization. All tags successfully transmitted data with deployment durations ranging from 6 to 124 days. Tag data revealed shark movements in excess of 200 km from initial tagging locations, with sharks primarily utilizing offshore waters associated with the continental shelf edge from Desoto Canyon to the Texas/Mexican border. While most sharks remained in US waters, one individual moved from the northern Gulf of Mexico into the Bay of Campeche off the coast of Mexico. Sharks spent 87 % of their time between 20 and 125 m and 83 % of their time in waters between 23 and 30 °C. Since dusky sharks are among the most vulnerable shark species to fishing mortality, there is a recovery plan in place for US waters; however, since they have been shown to make long-distance migrations, a multi-national management plan within the WNA may be needed to ensure the successful recovery of this population.     

Shifting elasmobranch community assemblage at Cocos Island—an isolated marine protected area

Fishing pressure has increased the extinction risk of many elasmobranch (shark and ray) species. Although many countries have established no‐take marine reserves, a paucity of monitoring data means it is still unclear if reserves are effectively protecting these species. We examined data collected by a small group of divers over the past 21 years at one of the world's oldest marine protected areas (MPAs), Cocos Island National Park, Costa Rica. We used mixed effects models to determine trends in relative abundance, or probability of occurrence, of 12 monitored elasmobranch species while accounting for variation among observers and from abiotic factors. Eight of 12 species declined significantly over the past 2 decades. We documented decreases in relative abundance for 6 species, including the iconic scalloped hammerhead shark (Sphyrna lewini) (?45%), whitetip reef shark (Triaenodon obesus) (?77%), mobula ray (Mobula spp.) (?78%), and manta ray (Manta birostris) (?89%), and decreases in the probability of occurrence for 2 other species. Several of these species have small home ranges and should be better protected by an MPA, which underscores the notion that declines of marine megafauna will continue unabated in MPAs unless there is adequate enforcement effort to control fishing. In addition, probability of occurrence at Cocos Island of tiger (Galeocerdo cuvier), Galapagos (Carcharhinus galapagensis), blacktip (Carcharhinus limbatus), and whale (Rhincodon typus) sharks increased significantly. The effectiveness of MPAs cannot be evaluated by examining single species because population responses can vary depending on life history traits and vulnerability to fishing pressure.     

Demographic Processes Underlying Population Growth and Decline in Salamandra salamandra

Abstract:  Human activity commonly has negative impacts on wildlife. Often, however, only a single element of the life cycle is affected, and it is unclear whether such effects translate into effects on population growth. This is particularly true for research into the causes of global amphibian declines, where experimental research focuses primarily on the aquatic larval stages but theory suggests these stages have only minor importance for population growth. We used data from long-term mark-recapture studies of two natural populations of the salamander Salamandra salamandra to confirm the predictions of population models. One population remained stable (i.e., stationary) throughout the 20 years of the study whereas the other declined to local extinction. We used mark-recapture models to break down population growth rate into its two main components, recruitment and adult survival. Survival of postmetamorphic salamanders was constant over time in the stable population, whereas the declining population was characterized by a decrease in survival and constant recruitment. Population growth was most sensitive to variation in adult survival. Current amphibian research focuses on preadult stages, and researchers assume recruitment is the most important determinant of population growth. This may not be the case. A better understanding of amphibian population dynamics is possible only through the integration of experiments, theory, and data from natural populations. Our results also suggest that amphibian conservation efforts should focus on all stages of the life cycle and their associated habitats.     

Population Trends in Pacific Oceanic Sharks and the Utility of Regulations on Shark Finning

Accurate assessment of shark population status is essential for conservation but is often constrained by limited and unreliable data. To provide a basis for improved management of shark resources, we analyzed a long‐term record of species‐specific catches, sizes, and sexes of sharks collected by onboard observers in the western and central Pacific Ocean from 1995 to 2010. Using generalized linear models, we estimated population‐status indicators on the basis of catch rate and biological indicators of fishing pressure on the basis of median size to identify trends for blue (Prionace glauca), mako (Isurus spp.), oceanic whitetip (Carcharhinus longimanus), and silky (Carcharhinus falciformis) sharks. Standardized catch rates of longline fleets declined significantly for blue sharks in the North Pacific (by 5% per year [CI 2% to 8%]), for mako sharks in the North Pacific (by 7% per year [CI 3% to 11%]), and for oceanic whitetip sharks in tropical waters (by 17% per year [CI 14% to 20%]). Median lengths of silky and oceanic whitetip sharks decreased significantly in their core habitat, and almost all sampled silky sharks were immature. Our results are consistent with results of analyses of similar data sets. Combined, these results and evidence of targeted fishing for sharks in some regional fisheries heighten concerns for sustainable utilization, particularly for oceanic whitetip and North Pacific blue sharks. Regional regulations that prohibit shark finning (removal of fins and discarding of the carcass) were enacted in 2007 and are in many cases the only form of control on shark catches. However, there is little evidence of a reduction of finning in longline fisheries. In addition, silky and oceanic whitetip sharks are more frequently retained than finned, which suggests that even full implementation of and adherence to a finning prohibition may not substantially reduce mortality rates for these species. We argue that finning prohibitions divert attention from assessing whether catch levels are sustainable and that the need for management of sharks should not be addressed by measures that are simple to implement but complex to enforce and evaluate. Tendencias Poblacionales de Tiburones del Océano Pacífico y la Utilidad de Regulaciones sobre Cercenamiento de Aletas