Recent U.S. legislation applies a precautionary approach to setting catch regulations in federal fisheries management. A transparent approach to complying with federal guidelines involves calculating the catch recommendation that corresponds to a specified probability, P*, of exceeding the "true" overfishing limit (OFL) located within an estimated distribution. The P* methodology aims to manage the risk of overfishing explicitly, but choice of P* alone does not provide sufficient information on all of the risks associated with a control rule—both the probability of overfishing and the severity of overfishing. Rather, the ramifications of P* choices depend on the amount of uncertainty in the stock assessment and on the life history of the species in question. To evaluate these effects on the risks associated with P* rules, my study simulated fishing three example species under three levels of uncertainty. Trends identified among example species were consistent with predictions from life history. Periodic strategists, which have highly variable recruitment, experienced probabilities of overfishing which exceeded P* and which increased in time. Equilibrium strategists showed more predictable risks of overfishing but may have less capacity to recover from depleted biomass levels. Differences in the size of the OFL distribution—representing differences in levels of uncertainty—led to mixed results depending on whether the distribution was biased or whether uncertainty was fully characterized. Lastly, because OFL distributions are themselves estimates and subject to uncertainty in their shape and size, lower P* values closer to the tails of the estimated distribution produced more variable resulting risks. / Master of Science

Estimation of natural mortality within statistical catch-at-age models has been relatively unsuccessful and is uncommon within stock assessments. The models I created estimated population-dynamics parameters, including natural mortality, through Metropolis-Hastings algorithms from Gulf of Mexico red-snapper Lutjanus campechanus data. I investigated the influences of assumptions regarding model configuration of natural mortality and selectivity-at-age parameters by comparing multiple models. The results of this study are preliminary due to parameter estimates being bounded by uniform priors and thus a potential lack of convergence to the posterior distribution. Estimation of a natural-mortality parameter at age 0 or a Lorenzen natural-mortality parameter could be confounded with selectivity-at-age-1 parameters for bycatch from the shrimp fisheries. The Lorenzen natural-mortality curve was calculated by dividing the parameter by red snapper length at age. An age-1 natural-mortality parameter might not be estimable with the currently available data. Values of the natural-mortality parameter for ages 2 and older appear to be slightly less influenced by assumptions regarding selectivity-at-age parameters. We conducted a simulation study to determine the accuracy and precision of natural-mortality estimation assuming the selectivity-at-age-1 parameter for bycatch from the shrimp fisheries equaled 1.0 and a Lorenzen natural-mortality curve. The simulation study indicated that initial abundance-at-age parameters may be inestimable within the current model and may influence other parameter estimates. The preliminary simulation results showed that the Lorenzen natural-mortality parameter was consistently slightly underestimated and apical-fishing-mortality parameters were considerably underestimated. The estimation of natural mortality within a statistical catch-at-age model for Gulf of Mexico red snapper has many caveats and requires additional investigation. / Master of Science

Three hammerhead sharks (Sphyrna spp.) are currently managed as part of the large coastal shark complex in the United States. Including multiple species in an assessment ignores the different stock dynamics of each individual species within the complex due to different life histories. This study completed individual assessments of scalloped (S. lewini), great (S. mokarran), and smooth (S. zygaena) hammerhead sharks in the U.S. Atlantic Ocean and Gulf of Mexico. Combined data for all three species and unclassified hammerhead sharks were also used to produce a stock assessment of the hammerhead shark complex. Depletions of 83%, 96%, and 91% were estimated for scalloped, great, and smooth hammerhead sharks, respectively, between 1981 and 2005. When modeled as a single stock, the hammerhead shark complex experienced a 90% decline over the same time period. All three stocks, and the complex were overfished (below population size associated with maximum sustainable yield (MSY)), and overfishing (fishing level above that associated with MSY) occurred in 2005. We found that scalloped hammerhead shark population recovery is likely to occur within 10 years if catch remains at or below 2005 levels. Great and smooth hammerhead sharks will likely still be overfished in 30 years unless catches are reduced. It appears that the species composition could be changing in this hammerhead shark complex. The faster-growing scalloped hammerhead sharks are able to withstand fishing pressure better than great or smooth hammerhead sharks. However, it is difficult to target any single large coastal shark species while fishing; hence they are subject to similar fishing pressure. The result is a greater decline in great and smooth hammerhead sharks than experienced by scalloped hammerhead sharks. Therefore, the proportion of scalloped hammerhead sharks increased between 1981 and 2005. Species-specific stock assessments, such as those presented here, allow managers to more closely monitor populations of slower-growing species and reduce the risk of overexploitation of those species. / Master of Science

Chrysoblephus puniceus is an over-exploited linefish species, endemic to the coastlines off southern Mozambique and eastern South Africa. Over-exploitation and habitat loss are two of the biggest threats to the sustainability of fisheries globally. Assessing the genetic stock structure (a prerequisite for effective management) and predicting climate related range changes will provide a better understanding of these threats to C. puniceus which can be used to improve the sustainability of the fishery. Two hundred and eighty four genetic samples were collected from eight sampling sites between Ponta da Barra in Mozambique and Coffee Bay in South Africa. The mitochondrial control region and ten microsatellite loci were amplified to analyse the stock structure of C. puniceus. The majority of microsatellite and mtDNA pairwise population comparisons were not significant (P > 0.05) although Xai Xai and Inhaca populations had some significant population comparisons for mtDNA (P < 0.05). AMOVA did not explain any significant variation at the between groups hierarchical level for any pre-defined groupings except for a mtDNA grouping which separated out Xai Xai and Inhaca from other sampling sites. SAMOVA, isolation by distance tests, structure analysis, principle component analysis and spatial autocorrelation analysis all indicated a single population of C. puniceus as being most likely. The migrate-n analysis provided evidence of current driven larval transport, with net migration rates influenced by current dynamics.Two hundred and thirty six unique presence points of C. puniceus were correlated with seasonal maximum and minimum temperature data and bathymetry to model the current distribution and predict future distribution changes of the species up until 2030. Eight individual species distribution models were developed and combined into a mean ensemble model using the Biomod2 package. Winter minimum temperature was the most important variable in determining models outputs. Overall the ensemble model was accurate with a true skills statistic score of 0.962. Binary transformed mean ensemble models predicted a northern and southern range contraction of C. puniceus' distribution of 15 percent; by 2030. The mean ensemble probability of occurrence models indicated that C. puniceus' abundance is likely to decrease off the southern Mozambique coastline but remain high off KwaZulu-Natal. The results of the genetic analysis support the theory of external recruitment sustaining the KwaZulu Natal fishery for C. puniceus. While the high genetic diversity and connectivity may make C. puniceus more resilient to disturbances, the loss of 15 percent; distribution and 11 percent; genetic diversity by 2030 will increase the species vulnerability. The decrease in abundance of C. puniceus off southern Mozambique together with current widespread exploitation levels could result in the collapse of the fishery. A single transboundary stock of C. puniceus highlights the need for co-management of the species. A combined stock assessment between South Africa and Mozambique and the development of further Marine Protected Areas off southern Mozambique are suggested as management options to minimise the vulnerability of this species.

The study of spatiotemporal dynamics of fish populations is important for both stock assessment and fishery management. I explored the impacts of environmental and anthropogenic factors on spatiotemporal patterns of fish populations, and contributed to stock assessment and management by incorporating the inherent spatial structure. Hierarchical models were developed to specify spatial and temporal variations, and Bayesian methods were adopted to fit the models. Yellow perch (Perca flavescens) is one of the most important commercial and recreational fisheries in Lake Erie, which is currently managed using four management units (MUs), with each assessed by a spatially-independent stock-specific assessment model. The current spatially-independent stock-specific assessment assumes that movement of yellow perch among MUs in Lake Erie is statistically negligible and biologically insignificant. I investigated whether the assumption is violated and the effect this assumption has on assessment. I first explored the spatiotemporal patterns of yellow perch abundance in Lake Erie based on data from a 27-year gillnet survey, and analyzed the impacts of environmental factors on spatiotemporal dynamics of the population. I found that yellow perch relative biomass index displayed clear temporal variation and spatial heterogeneity, however the two middle MUs displayed spatial similarities. I then developed a state-space model based on a 7-year tag-recovery data to explore movements of yellow perch among MUs, and performed a simulation analysis to evaluate the impacts of sample size on movement estimates. The results suggested substantial movement between the two stocks in the central basin, and the accuracy and precision of movement estimates increased with increasing sample size. These results demonstrate that the assumption on movements among MUs is violated, and it is necessary to incorporate regional connectivity into stock assessment. I thus developed a tag-integrated multi-region model to incorporate movements into a spatial stock assessment by integrating the tag-recovery data with 45-years of fisheries data. I then compared population projections such as recruitment and abundance derived from the tag-integrated multi-region model and the current spatial-independent stock-specific assessment model to detect the influence of hypotheses on with/without movements among MUs. Differences between the population projections from the two models suggested that the integration of regional stock dynamics has significant influence on stock estimates. American Shad (Alosa sapidissima), Hickory Shad (A. mediocris) and river herrings, including Alewife (A. pseudoharengus) and Blueback Herring (A. aestivalis), are anadromous pelagic fishes that spend most of the annual cycle at sea and enter coastal rivers in spring to spawn. Alosa fisheries were once one of the most valuable along the Atlantic coast, but have declined in recent decades due to pollution, overfishing and dam construction. Management actions have been implemented to restore the populations, and stocks in different river systems have displayed different recovery trends. I developed a Bayesian hierarchical spatiotemporal model to identify the population trends of these species among rivers in the Chesapeake Bay basin and to identify environmental and anthropogenic factors influencing their distribution and abundance. The results demonstrated river-specific heterogeneity of the spatiotemporal dynamics of these species and indicated the river-specific impacts of multiple factors including water temperature, river flow, chlorophyll a concentration and total phosphorus concentration on their population dynamics. Given the importance of these two case studies, analyses to diagnose the factors influencing population dynamics and to develop models to consider spatial complexity are highly valuable to practical fisheries management. Models incorporating spatiotemporal variation describe population dynamics more accurately, improve the accuracy of stock assessments, and would provide better recommendations for management purposes. / Doctor of Philosophy / Many fish populations exhibit complex spatial structure, but the spatial patterns have been incorporated into stock assessment only in few cases. A full understanding of spatial structure of fish populations is needed to better manage the populations. Stock assessment and management strategies should depend on the inherent spatial structure of the target fish population. There have been many approaches developed to analyze spatial structure of fish populations. In this dissertation, I developed quantitative models to analyze fish demographic data and tagging data to explore spatial structure of fish populations. Yellow perch (Perca flavescens) in Lake Erie and Alosa group including American Shad (Alosa sapidissima), Hickory Shad (A. mediocris) and river herrings (Alewife A. pseudoharengus and Blueback Herring A. aestivalis) in selected tributaries of the Chesapeake Bay were taken as examples. Fishery-independent data for yellow perch displayed spatial similarities in the central basin of Lake Erie. Distinct temporal trends were observed in relative abundance data for Alosa sp. in different tributaries of the Chesapeake Bay. Substantial yellow perch movement among the central basin of the Lake was observed in tagging data. Ignoring the inherent spatial structure may cause fish to be overfished in some regions and underfished in others. To maximize the effectiveness of management in all regions for fish populations, I highly recommend incorporating spatial structure into stock assessment and management such as the ones developed in this dissertation.

The catchability coefficient is used in most marine stock assessment models, and is usually assumed to be stationary and density-independent. However, recent research has shown that these assumptions are violated in most fisheries. Violation of these assumptions will cause underestimation of stock declines or recoveries, leading to inappropriate management policies. This project assesses the soundness of stationarity and density independence assumptions using multi-species data for seven stocks and four gears in the U.S. Gulf of Mexico. This study also develops a multi-species methodology to compensate for failures of either assumption. To evaluate catchability assumptions, abundance-at-age was reconstructed and compared with catch-per-unit-effort data in the Gulf. Mixed-effects, Monte Carlo, and bootstrap analyses were applied to estimate time-varying catchability parameters. Gulf data showed large and significant density dependence (0.71, s.e. 0.07, p<0.001) and increasing trends in catchability (2.0% annually compounding, s.e. 0.6%, p < 0.001). Simulation modeling was also used to evaluate the accuracy and precision of seven different single-species and multi-species estimation procedures. Imputing estimates from similar species provided accurate estimates of catchability parameters. Multi-species estimates also improved catchability estimation when compared with the current assumptions of density independence and stationarity. This study shows that multi-species data in the Gulf of Mexico have sufficient quantity and quality to accurately estimate catchability model parameters. This study also emphasizes the importance of estimating density-dependent and density-independent factors simultaneously. Finally, this study shows that multi-species imputation of catchability estimates decreases errors compared with current assumptions, when applied to single-species stock assessment data. / Master of Science

Poenskop Cymatoceps nasutus (Pisces: Sparidae), an endemic South African sparid, is an important angling species being predominantly targeted by the recreational shore and skiboat sector. This species is slow-growing, long-lived, late-maturing and sex-changing, making poenskop acutely sensitive to over-exploitation. Despite interventions, such as the imposition of size and bag limits (currently 50 cm TL and one per licensed fisher per day) by authorities, catch-per-unit-effort trends reflect a severe and consistent stock decline over the last two decades. Poenskop has been identified as a priority species for research and conservation. Although the biology and population dynamics of this species have been well-documented, little is known about the movement behaviour of poenskop. Furthermore, there is a complete lack of information on its genetic stock structure. This thesis aimed to address the current knowledge gaps concerning movement behaviour and genetic stock structure of poenskop, making use of a range of methods and drawing on available information, including available fishery records as well as published and unpublished survey and research data, and data from long-term monitoring programmes. Analysis of available catch data (published and unpublished) revealed a decline in the number of poenskop caught as well as size of fish taken over the last two decades, ultimately reflecting the collapse of the stock (estimated to be at 20% of their pristine level). Improved catch-per-unit-effort data from the Tsitsikamma National Park Marine Protected Area (MPA), and larger poenskop being caught in the no-take areas than adjacent exploited areas of the Pondoland MPA confirmed that MPAs can be effective for the protection and management of poenskop. The current MPA network in South Africa is already wellestablished, and encompasses considerable reef areas, being preferable for poenskop habitation. Conventional dart tagging and recapture information from three ongoing, long-term fishtagging projects, conducted throughout the poenskop’s distribution, indicated high levels of residency at all life-history stages. Coastal region, seasonality and time at recapture did not appear to have a significant effect on the level of movement or distance moved. However, on examining the relationship among coastal movements and fish size and ages, larger and older fish (adults) moved greater distances, with juveniles and sub-adults showing high degrees of residency. An estimation of home-range size indicated smaller poenskop to hold smaller home-ranges, while larger poenskop hold larger home-ranges. Large easterly displacements of a number of adult poenskop is in accordance with previous findings that this species may undertake a unidirectional migration up the coastline of South Africa where they possibly settle in Transkei waters for the remainder of their lives. This high level of residency makes poenskop vulnerable to localised depletion, although they can be effectively protected by suitable MPAs. Despite considerable tagging effort along the South African coastline (2 704 poenskop tagged with 189 recaptures, between 1984 and 2010), there remains limited information on the connectivity of different regions along the South African coastline. This was addressed using mitochondrial DNA sequencing. The mitochondrial DNA control region was used due to its high substitution rate, haploid nature, maternal inheritance and absence of recombination. The mtDNA sequencing showed no evidence of major geographic barriers to gene flow in this species. Samples collected throughout the core distribution of poenskop showed high genetic diversity (h = 0.88, π = 0.01), low genetic differentiation among regions, no spatial structure (ɸST = 0.012, p = 0.208) and no evidence of isolation by distance. The collapsed stock status of poenskop as well as the fact that it is being actively targeted by recreational and commercial fishers suggests that this species requires improved management, with consideration given to its life-history style, residency and poor conservation status. Management recommendations for poenskop, combined with increasing South Africa’s existing MPA network, include the possibility of setting up a closed season (during known spawning periods) as well as the decommercialisation of this species. The techniques used and developed in this study can also be adopted for other overexploited linefish species.

Marine Protected Areas (MPAs), the Ecosystem Approach to Fisheries management (EAF) and Integrated Coastal Management (ICM) have been identified as possible alternatives to traditional linefish management measures, which have largely failed. Monitoring and assessment of fish communities on a long-term basis is necessary, and will provide a means to evaluate the effectiveness of such management measures. Therefore, standardised protocols and optimal sampling methods for long-term monitoring (LTM) and assessment of coastal fish communities are essential. This study aimed to identify suitable methods and develop a protocol for assessment of inshore reef fish communities. A suitable location for evaluation of proposed methods was identified in the warm temperate biogeographical region of South Africa, encompassing the well-established Tsitsikamma Coastal National Park MPA and an adjacent exploited area. Chrysoblephus laticeps (roman) was identified as an indicator species for the study, as it has been well-studied and is well represented in the area. Underwater visual census (UVC) and controlled fishing were identified as suitable methods. UVC transects were found to be superior to point counts, in terms of sampling efficiency, variability, bias and required sample size. An effort of two angler hours per fishing station was shown to provide low catch variability, while at the same time a representative catch and low overall cost and required time. The methods were incorporated in a proposed sampling protocol, and evaluated. The methods were able to detect known differences between protected and exploited communities. It is recommended that LTM within protected areas, for detection of natural change, be focused on community-level indicators, while LTM in exploited areas, aimed at detection of anthropogenic change, be focused on species-level indicators. The proposed protocol with standardised methods will allow for comparisons across a network of LTM sites and provide the opportunity for a broad-scale assessment of the effects of environmental variables on reef fish stocks. The protocol developed in this study has application in other biogeographical regions in South Africa, and other parts of the world. Shift in the focus of much marine research, in South Africa and elsewhere, to LTM, highlights the relevance and timeous nature of this study.

It is currently hypothesised that the chokka squid (Loligo reynaudi) consist of a single stock. This was tested through a spatial comparison of the morphology of this species. Forty three morphometric characters were measured from 1079 chokka squid collected from three regions: the south coast of South Africa, the west coast of South Africa, and southern Angola. While no significant differences were found for the hard body parts, results from discriminant function analyses showed the soft body morphometric characters from each of the three regions differed, with an overall correct classification rate of 100% for males and 99% for females in all three regions. Due to the existing model being used to assess the resource currently being updated it was not feasible to apply this model to the area-disaggregated data from this study. Rather, the CPUE trends and catches from the area-disaggregated data were compared against those of the area-aggregated data, as a first attempt to discern any appreciable differences which would suggest the use of disaggregated data in future assessments. Both the trawl and jig CPUE trends from the area-disaggregated analysis differed only slightly from those of the area-aggregated data. Similarly, the spring and autumn biomass trends for the main spawning area (east of 22°E) followed the same trends as for the full area. It is therefore concluded that there is currently no need to assess the resource on an area-disaggregated basis.

White steenbras Lithognathus lithognathus (Pisces: Sparidae) has been a major target species of numerous fisheries in South Africa, since the late 19th century. Historically, it contributed substantially to annual catches in commercial net fisheries, and became dominant in recreational shore catches in the latter half of the 20th century. However, overexploitation in both sectors resulted in severe declines in abundance. The ultimate collapse of the stock by the end of the last century, and the failure of traditional management measures to protect the species indicate that a new management approach for this species is necessary. The species was identified as a priority for research, management and conservation in a National Linefish Status Report. Despite knowledge on aspects of its biology and life history, little is known about juvenile habitat use patterns, home range dynamics and movement behaviour in estuaries. Similarly, the movement and migration of larger juveniles and adults in the marine environment are poorly understood. Furthermore, there is a complete lack of information on its genetic stock structure. Such information is essential for effective management of a fishery species. This thesis aimed to address the gaps in the understanding of white steenbras movement patterns and genetic stock structure, and provide an assessment of its current conservation status. The study adopted a multidisciplinary approach, incorporating a range of methods and drawing on available information, including published literature, unpublished reports and data from long-term monitoring programmes. Acoustic telemetry, conducted in a range of estuaries, showed high site fidelity, restricted area use, small home ranges relative to the size of the estuary, and a high level of residency within estuaries at the early juvenile life stage. Behaviour within estuaries was dominated by station-keeping, superimposed by a strong diel behaviour, presumably based on feeding and/or predator avoidance, with individuals entering the shallow littoral zone at night to feed, and seeking refuge in the deeper channel areas during the daytime. Conventional dart tagging and recapture data from four ongoing, long-term coastal fish tagging projects, spread throughout the distribution of this species, indicated high levels of residency in the surf zone at the late juvenile and sub-adult life stages. Consequently, juvenile and sub-adult white steenbras are vulnerable to localised depletion, although they can be effectively protected by suitably positioned estuarine protected areas (EPAs) and marine protected areas (MPAs), respectively. It has been hypothesized that adult white steenbras undertake large-scale coastal migrations between summer aggregation areas and winter spawning grounds. The scale of observed coastal movements was correlated with fish size (and age), with larger fish undertaking considerably longer-distance coastal movements than smaller individuals, supporting this hypothesis. Given the migratory behaviour of adults, and indications that limited spawning habitat exists, MPAs designed to protect white steenbras during the adult life stage should encompass all known spawning aggregation sites. The fishery is plagued by problems such as low compliance and low enforcement capacity, and alternative management measures, such as seasonal closure, need to be evaluated. Despite considerable conventional dart tagging effort around the coastline (5 782 fish tagged) with 292 recaptures there remains a lack of empirical evidence of fish migrating long distances (> 600 km) between aggregation and spawning areas. This uncertainty in the level of connectivity among coastal regions was addressed using mitochondrial DNA sequencing and genotyping of microsatellite repeat loci in the nuclear genome, which showed no evidence of major geographic barriers to gene flow in this species. Samples collected throughout the white steenbras core distribution showed high genetic diversity, low genetic differentiation and no evidence of isolation by distance or localised spawning. Although historically dominant in several fisheries, analysis of long-term commercial and recreational catch data for white steenbras indicated considerable declines and ultimately stock collapse. Improved catch-per-unit-effort in two large MPAs subsequent to closure confirmed that MPAs can be effective for the protection of white steenbras. However, the current MPA network encompasses a low proportion of sandy shoreline, for which white steenbras exhibits an affinity. Many MPAs do not prohibit recreational shore angling, which currently accounts for the greatest proportion of the total annual catch. Furthermore, EPAs within the juvenile distribution protect a negligible proportion of the total available surface area of estuaries – habitat on which white steenbras is wholly dependent. Despite some evidence of recent increases in abundance in estuaries and the surf zone in certain areas, white steenbras meets the criteria for “Endangered” on the IUCN Red List of Threatened Species, and for “Protected species” status on the National Environmental Management: Biodiversity Act of South Africa. The species requires improved management, with consideration for its life-history style, estuarine dependency, surf zone residency, predictable spawning migrations and its poor conservation status. The multidisciplinary approach provides valuable information towards an improved scientific basis for the management of white steenbras and a framework for research that can be adopted for other overexploited, estuarine-associated coastal fishery species.