Although the marine environment has generally been considered to be less in need of attention from conservationists, politicians and managers than the terrestrial environment, there has been an increasing appreciation that there is little left in the marine environment that can be considered unaltered by humans. Conservation action is therefore urgently needed to protect marine populations, species and environments from anthropogenic pressures. Since overharvesting seems to be the main cause of the collapse in coastal ecosystems, areas need to be protected so that stocks can recover. However, the proclamation of such marine protected areas (MPAs) need to be well planned to ensure that species are well represented and that their biological requirements are met in the long term. While terrestrial conservation planners have started to incorporate biological, social and economic information into so-called systematic conservation plans, marine planners have been more hesitant to use these approaches due to the greater openness of the ocean (with the associated greater dispersal of marine organisms and propogules, lower levels of endemism, and lack of ownership) and the paucity of available information. The current study aimed to investigate the applicability of systematic approaches to the process of selecting sites for a network of potential replenishment zones for seabream species that could supplement protection already afforded by existing no-fishing reserves. Since many of these seabream species are popular targets in both the recreational and commercial linefishery sectors, they have been overexploited and are in need of additional protection. Information on these species and the diversity of their associated reefs were obtained through consultations with scientists, managers and resource-users (namely SCUBA divers, spearfishers as well as recreational and commercial fishers). These data were obtained during workshops where Participatory Rural Appraisal (PRA) scoring and mapping methods were applied. Although ordinal-scale, rather than interval-or ratio-scale data were obtained, these data could be used in an interactive selection process. Because these methods require relatively cheap materials and few personnel, they could be used to collect information where limited budgets are available. Quantitative targets for the protection of individual species were set on the basis of their status in the Marine Living Resource Act, since this reflected their different requirements for protection. A preliminary investigation indicated that the current no-fishing reserves effectively achieved most of these goals for seabream species. The selection of sites for the replenishment network was based on irreplaceability analyses, while decisions between sites with lower irreplaceability values were based on explicit design rules. A total of 26 sites were required to achieve targets for all species. These replenishment zones could potentially be established, managed and monitored by voluntary community groups. This study therefore indicated that systematic conservation planning can successfully be applied to the marine environment, while the involvement of stakeholders is crucial to ensure that conservation action is supported.

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.

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.

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.

Η οικολογία και η δυναμική της εγκατάστασης εξετάστηκαν για τέσσερα μεσογειακά παράκτια είδη: Diplodus vulgaris, Diplodus sargus, Oblada melanura και Diplodus annularis. Η διερεύνηση τους έγινε με υποδιαίρεση της βενθικής ιχθυονυμφικής τους φάσης σε έξι οντογενετικά στάδια βάσει χρωματικών πρότυπων που εμφάνιζαν διαδοχή και ήταν χαρακτηριστικά για κάθε είδος. Η συλλογή δεδομένων έγινε για κάθε οντογενετικό στάδιο με στρωματοποιημένη δειγματοληψία, με τις στρώσεις να αποτελούν συνδυασμό τύπου υποστρώματος και βάθους στην παράκτια περιοχή ανατολικά των εκβολών του Αχελώου, από βάθος 0 έως 5m. Τα δεδομένα συλλέχθηκαν με απ’ ευθείας παρατηρήσεις με χρήση καταδυτικής συσκευής. Υπολογίστηκαν οι πυκνότητες κάθε οντογενετικού σταδίου στους διάφορους τύπους υποστρώματος και βάθη μέσα στο έτος. Η κατανομή των πυκνοτήτων αυτών έδωσε στοιχεία για τις οικολογικές προτιμήσεις των βενθικών ιχθυονυμφών και των νεαρών, καθώς και για τους ρυθμούς μεταμόρφωσης κάθε είδους. Επίσης ελέγχθηκε το πρότυπο συμπεριφοράς των ιχθυονυμφών του μελανουριού κατά την εγκατάσταση τους, μ’ επινόηση ειδικής μεθοδολογίας κι ενός δείκτη που απέδωσε ποσοτικά τις διάφορες εκφάνσεις της συμπεριφοράς. Σύμφωνα με τ’ αποτελέσματα της εργασίας, φάνηκε ότι ο σαργόπαπας και ο σαργός εγκαθίστανται την κρύα περίοδο του έτους, ενώ το μελανούρι και ο σπάρος τη θερμή. Όλα τα είδη εμφάνισαν γρηγορότερο ρυθμό μεταμόρφωσης κατά τα πρώτα στάδια, ενώ τα είδη που εγκαταστάθηκαν το καλοκαίρι είχαν συνολικά γρηγορότερους ρυθμούς μεταμόρφωσης από τα χειμερινά. Τα είδη εμφάνισαν διαφοροποίηση στη χρονική και χωρική κατανομή τους ώστε να ελαττώνονται οι ανταγωνιστικές δράσεις. Το είδος με τη μακρύτερη χρονική παρουσία ήταν ο σαργόπαπας (10 μήνες), ο οποίος εμφάνισε ευρύτητα ως προς τις οικολογικές του απαιτήσεις διασπειρόμενος σ’ όλο το εύρος βάθους και τύπους υποστρώματος, εκτός του αμμώδους. Στον αντίποδα, το μελανούρι είχε τη μικρότερη συνολική παρουσία (4 μήνες) και απόλυτη εξάρτηση από σκληρό υπόστρωμα. Ο σαργός κι ο σπάρος έδειξαν ενδιάμεσες οικολογικές απαιτήσεις με προτίμηση στα σκληρά υποστρώματα και στα φανερόγαμα αντίστοιχα. Κανένα είδος δεν έδειξε προτίμηση σε αμμώδεις βυθούς. Ιδιαίτερο οικολογικό ρόλο εμφάνισε το ρηχότερο τμήμα της παράκτιας ζώνης και οι μεταβατικές ζώνες διαφορετικών υποστρωμάτων. Οι βενθικές ιχθυονύμφες του μελανουριού επέδειξαν τάση παραμονής για μέρες με μικρές ομάδες σε σημεία του υποστρώματος. Στα επόμενα στάδια σχημάτισαν μεγάλες ομάδες με μικρότερη εξάρτηση από το υπόστρωμα. / The ecology and dynamics of settlement process were studied for four sparids (Diplodus vulgaris, D. sargus. D. annularis and Oblada melanura), in a coastal littoral in western Hellas. The benthic larval phase of each species was divided in six ontogenetic stages, according to specific patterns in appearance that followed a sequence. Preliminary observations showed non homogeneous distribution of the larvae in the area, so stratified sampling was applied. Strata represented the different substratum types accounted in each of the three depth zones of 0-1m, 1-2m and 2-5m. Data concerned abundances of each ontogenetic stage in each stratum during a 15 month interval and they were collected by visual census. Moreover, data on site-attachment ontogenetically depended behavior of benthic larvae of O. melanura were also collected. The later behavior was studied via an index devised for quantifying different expressions of residence behavior. Two of the studied species (D. vulgaris and D. sargus), settled during the cold period of the year while the other two settled during summer. Metamorphosis rates were faster at the first ontogenetic stages for all species, while they were faster in total for the summer species. Temporal and spatial distribution of the species was arranged in order to minimize competition. D. vulgaris and D. sargus settled in the same substratum types in different periods and O. melanura and D. annularis settled during the same period in different substrata. D. vulgaris remained in the nursery area the longest period, exhibiting the broadest ecological requirements, being distributed in different substrata and depths. O. melanura remained half as long in the nursery area, exhibiting the narrowest ecological requirements, staying always over hard substratum in shallow waters. The other two species exhibited intermediate conditions, with D. sargus preferring hard substratum and D. annularis, seagrass beds. None of the studied species preferred soft substratum. The shallowest part of the littoral appeared to be very important for three species (D. vulgaris, D. sargus, O. melanura) and the transition zones (between two different substrata), for all four of them. Benthic larvae of Oblada melanura commenced settlement in small shoals that resided for varying number of days in specific sites over stones or rocky substratum. Gradually as metamorphosis proceeded they became more kinetic forming larger shoals that expanded their home range.

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.