The zebra sea bream, Diplodus cervinus (Sparidae) is an inshore fish comprised of two boreal subspecies from the Gulf of Oman and the Mediterranean / north eastern Atlantic and one austral subspecies from South Africa and southern Angola. The assumption of a single austral subspecies has, however, been questioned due to mounting molecular and morphological evidence suggesting that the cool Benguela current is a vicariant barrier that has separated many synonymous inshore fish species between South Africa and southern Angola. The aims of this thesis are to conduct a comparative morphological analysis of Diplodus cervinus in southern Angola and South Africa in order to classify the southern Angolan population and then to conduct a life history assessment to assess the life history impact of allopatry on this species between the two regions. Results of the morphological findings of the present study (ANOSIM, p < 0.05, Rmeristic = 0.42) and (Rmorphometric = 0.30) along with a concurrent molecular study (FST = 0.4 – 0.6), identified significant divergence between specimens from South Africa (n = 25) and southern Angola (n = 37) and supported stock separation and possibly sub-speciation, depending on the classification criteria utilised. While samples from the two boreal subspecies were not available for the comparative morphological or molecular analysis, comparisons of the colouration patterns between the three subspecies, suggested similarity between the southern Angola and the northern Atlantic / Mediterranean populations. In contrast, the colouration patterns between the southern Angolan and South African specimens differed substantially, further supporting the morphological and molecular results. The distinct morphological divergence between the southern Angolan and South African populations was not reflected within the life history traits of both populations. A combination of methods, including length/age frequency analyses, adult sex ratios and histological analysis was used to determine that this species is a rudimentary hermaphrodite in southern Angola. Peak spawning season was observed between June and July. The overall sex ratio (M: F) was 1:1.52 with females dominating smaller younger size classes and 50% maturity was attained at 210 mm FL and 4.6 years. Females [L(t) = 287.5(1-e⁻°·¹⁸⁽t⁻²·⁸⁴⁾)] grew significantly faster (LRT, p < 0.05) than males [L(t) = 380.19(1-e⁻°·°⁶⁽t⁻⁷·¹²⁾)]. The higher maximum age of the southern Angolan population of D. cervinus (43 years) was older than that of South African individuals sampled in the tsitsikamma national park. The similarities in the life history of the two austral populations are probably a consequence of similar selective pressures in the similar warmtemperate habitats. Evidence to support the above comments was found in the feeding study which showed that the South African and Angolan populations were almost identical, with both populations feeding primarily on amphipods and polychaete worms throughout ontogeny. In contrast, the diet of their boreal conspecifics from the Mediterranean was different, where larger individuals tended to select larger, and more robust, prey items. The life history differences observed between the boreal and austral populations can be attributed to either sampling bias or environmental factors. Sampling biases included the use of different age and growth estimation techniques, while the environmental factors would include differential selective pressures most likely driven by different resource availability and exploitation. The present study provides crucial baseline life history information of a potentially exploitable species off southern Angola as well as information on the life history plasticity of the species. Unfortunately, the current lack of uniformity in the methods used to estimate life history parameters between studies conducted on the boreal and austral populations have complicated our understanding of the evolution of various life history trends in sparid fish. From a management perspective however, the results from the present study can be used to propose management strategies for an emerging trap fishery in southern Angola. Using a balanced exploitation fishery approach (harvesting up to the size-at-100% maturity), the size of the fish traps entrance was calculated based the morphological information from this and other small sparid species that are targeted and was estimated to be 62 mm.

The blacktail, Diplodus capensis, is an inshore sparid fish distributed from Mozambique to Angola. This species forms an important component of coastal fisheries within its distribution, one being the subsistence handline fishery in southern Angola. With this fishery being critically important to the livelihoods of local communities, a biological study and stock assessment was conducted to provide information for the management of this species in southern Angola. However, with molecular evidence suggesting that the Benguela current may have separated the southern African populations of many inshore fish species over two million years ago, a morphological, taxonomic analysis was considered necessary to first investigate whether there was evidence for allopatry in this species. A total of 46 morphometric measurements and 18 counts were carried out on specimens collected from various locations in southern Angola and South Africa. Results were analysed using multi-dimensional scaling (MDS) and the significance of clusters was tested using analysis of similarities (ANOSIM). Biological samples of D. capensis were collected monthly from an unexploited area from April 2008 to March 2009. Additional biological samples were collected from the subsistence fishers in an exploited area during May, June and December 2009. Standard biological laboratory techniques were employed for the lifehistory comparison between the exploited and unexploited area. A per-recruit analysis was conducted using the life-history parameters from both areas in order to assess the current status of the subsistence fishery and to investigate the potential short-falls of the per-recruit assessment approach. The morphometric comparison showed that there was not sufficient evidence for speciation between the southern Angolan and South African populations of D. capensis. There was, however, sufficient morphological evidence to suggest that these populations are separate stocks. This indicated that the existing reference points on which the management of the South African population is based are unsuitable for the Angolan population. Diplodus capensis in southern Angola is omnivorous, feeding predominantly on algae, barnacles and mussels. An ontogenetic shift from algae to barnacles and mussels was correlated with allometric growth patterns in their feeding apparatus. This species is a rudimentary hermaphrodite in southern Angola with peak spawning in June and July. The overall sex ratio (M: F) was 1: 4.7 in the unexploited area and 50% maturity was attained at 149.5mm FL and five years. Diplodus capensis in southern Angola exhibits very slow growth with the maximum age observed being 31 years (validated using mark recapture of chemically injected fish). Females [L(t) = 419.5(1-e⁻°·°⁴⁵⁽t⁻³·⁴ ⁾)] grew significantly faster (LRT, p < 0.05) than males [L(t) = 297.4(1-e⁻°·°⁷⁷⁽t⁻²·⁷⁾)], and females dominated the larger size classes and older age classes. In the exploited area, the length and age frequencies were severely truncated, the maximum observed age was greatly reduced (17 years) and the sex ratio was less female biased at 1: 2.2. Although there was no evidence for a physiological response to exploitation through alterations in growth or size/age at sexual maturity between the two areas, there was an increase in the proportion of small females in the exploited area, which may have been a compensatory response for the loss of large females. A combination of an underestimate of longevity, different estimates of the Von Bertalanffy growth parameters and overestimates of the natural mortality rate in the exploited population resulted in a 92% underestimate of the pristine spawner biomass-per-recruit (SBR) value. An assessment based on the actual pristine SBR estimate from the unexploited area revealed that the subsistence fishery had actually reduced D. capensis to 20% of its pristine SBR levels and highlighted the value of pre-exploitation life-history information for the application of per-recruit models. This study has shown that D. capensis in southern Angola displays life-history characteristics that render it susceptible to overexploitation, even at low levels of fishing pressure. The current lack of infrastructure and enforcement capacity in the fisheries department of Angola renders traditional linefish regulatory tools, such as size limits, bag limits and closed seasons, inappropriate. Therefore, suitably designed marine protected areas are recommended as the best management option for this species.

Overfishing in the Benguela Current Large Marine Ecosystem (BCLME) resulted in degradation of the ecosystem. This study used the Driver-Pressure-State-Impact-Response (DPSIR) indicator framework to determine whether the ecosystem is now recovering. Indicator trends were analysed using various data sources that included government institutions and intergovernmental institutions. The results showed that the overall effect of Driver indicators was negative. This was mainly because of socio-economic pressure such as the need to create more jobs in light of rising national unemployment and the declining contribution of the fisheries sector to Gross Domestic Product (GDP). In addition to scientific advice, socio-economic factors also influenced the determination of Total Allowable Catches (TACs). The overall trend of Pressure indicators was positively influenced by the effect of TACs. The TACs reduced the quotas allocated for commercial fishing. Environmental factors did not seem to play a significant role in this study. State indicators had mixed results with the indicators assessed almost split in the middle between those showing a positive trend and those showing a negative trend. On the other hand, Impact and Response indicators showed overall positive results. Therefore, the conclusion of the study was that the degradation of the BCLME has slowed down and there are some signs of recovery. / Environmental Sciences / M.Sc. (Environmental Science)

Overfishing in the Benguela Current Large Marine Ecosystem (BCLME) resulted in degradation of the ecosystem. This study used the Driver-Pressure-State-Impact-Response (DPSIR) indicator framework to determine whether the ecosystem is now recovering. Indicator trends were analysed using various data sources that included government institutions and intergovernmental institutions. The results showed that the overall effect of Driver indicators was negative. This was mainly because of socio-economic pressure such as the need to create more jobs in light of rising national unemployment and the declining contribution of the fisheries sector to Gross Domestic Product (GDP). In addition to scientific advice, socio-economic factors also influenced the determination of Total Allowable Catches (TACs). The overall trend of Pressure indicators was positively influenced by the effect of TACs. The TACs reduced the quotas allocated for commercial fishing. Environmental factors did not seem to play a significant role in this study. State indicators had mixed results with the indicators assessed almost split in the middle between those showing a positive trend and those showing a negative trend. On the other hand, Impact and Response indicators showed overall positive results. Therefore, the conclusion of the study was that the degradation of the BCLME has slowed down and there are some signs of recovery. / Environmental Sciences / M.Sc. (Environmental Science)