The aims of this study are to improve the understanding of the life cycle biology of L. forbesii and examine the ecological importance of this squid in the marine ecosystem. The life history study was based on two years of sampling during July 2006-June 2008 and, for the first time in this study area, made use of age data based on reading growth increments in statoliths. The statolith increment analysis suggests that the life-span of L. forbesii may be longer than one year, and hatching apparently occurred throughout the year. Two modes in size at maturity and a single mode in age at maturity suggest differences in growth rates between males and females as well as the existence of alternative growth strategies. The approach to statistical modeling was based on GAMs and revealed that growth and maturation of L. forbesii were influenced by both internal and external factors, in which their effect was sex-specific. This study provides evidence that males continue to invest energy for somatic growth, including growth in length, as they mature, while females invest more energy in growing reproductive tissue. A new study of squid diet, based on data from 2006-07 showed that clupeid fish were less important in the squid diet compared to a previous study in 1990-92, while the importance of gobies had increased. Modelling results revealed the influence of squid (predator) size, season and region on variation in diet and prey size of L. forbesii. The observed changes in the diet of L. forbesii over time may be related to changes in fish abundance. The results from the ecosystem model constructed using Ecopath with Ecosim suggest that squids have both direct negative impacts on their prey and indirect negative impact on other predators. The results from simulation suggest that increasing squid fishing intensity could impact most groups in the system, not only groups with direct trophic relationships with squid as prey or predators.

En el presente trabajo se evalúa la estructura histológica de las gónadas del calamar Loligo gahi como también su comportamiento reproductivo en determinados meses. De Enero a Marzo de 2007 se realizaron muestreos mensuales de 56 ejemplares de calamar Loligo gahi del muelle de la playa Pescadores de la costa de Lima, Perú. Fueron procesadas por técnica histológica convencional de hematoxilina–eosina para poder determinar las etapas de la ovogénesis, espermiogénesis y describir la organización histológica del ovario y del testículo. En las hembras de Loligo gahi la ovogénesis presenta 6 etapas o fases de desarrollo. En el macho la espermiogénesis constó de 5 etapas de desarrollo, lo que facilitará predecir adecuadamente el ciclo reproductivo. El análisis histológico se complementó con la obtención del índice gonadosomático, cuyos resultados permiten predecir que los meses de enero a marzo son los más adecuados para la maduración sexual y el desove en esta especie. The present work evaluates the histological structure of gonads of the squid Loligo gahi as well as their behavior in certain months. From January to March 2007 were conducted monthly sampling of 56 specimens of the squid Loligo gahi Beach pier fishermen along the coast of Lima, Peru. Were processed by conventional histological technique of hematoxylin-eosin to determine the stages of ovogénesis, espermiogénesis and describe the histological organization of the ovary and testis. In females of Loligo gahi 6 presents the ovogénesis phases or stages of development. In the male espermiogénesis consisted of 5 stages of development, which provide adequately predict the reproductive cycle. The histological analysis was supplemented by obtaining Gonadosomatic The results predict that the months of January through March are the most suitable for sexual maturation and spawning in this species

Thesis (MTech (Oceanography))Cape Peninsula University of Technology, 2010 / The current dynamics along the Tsitsikamma coast is described from a combination of acoustic current measurements. satellite-tracked surface drifters and underwater temperature recordings made between November 2006 and March 2008. The Tsitsikamma coast is largely a Marine Protected Area (MPA) that protects a rich marine biodiversity. The nearshore currents are important in the dispersal of eggs and larvae of many marine species. including the paralarvae of the commercially caught chokka squid. LoNgo reynaudii. Changes in the environment, including the currents. can affect the successful recruitment of chokka squid, and can bring about large annual fluctuations in biomass that creates economic uncertainty in the squid fishery. Results confirm the existence of a predominantly alongshore current off the Tsitsikamma coast. At Middelbank eastward flow was slightly dominant, with a percentage occurrence of 58% vs. 41% westward flow near the surface. The percentage eastward flow decreased with depth, with 41% vs. 58% westward flow near the seabed. At Thyspunt westward and eastward flow occurred at near equal percentages. but westward flow was slightly dominant throughout the water column. The alongshore current was strongest near the surface during eastward flow (maximum = 141 crn.s1: average = 27 crn.s '). while westward surface currents were weaker (maximum velocity = 78 cm.s1: average = 19 crn.s1). Current speed generally decreased with depth and opposing surface and bottom currents, associated with a thermal stratified water column, were occasionally recorded. The nearshore flow regime was characterised by frequent barotropic alongshore reversals that occurred year round. An increase in strong eastward episodes, and opposing surface and bottom currents during spring and summer months have implications for the dispersal of squid paralarvae during the summer and winter spawning seasons. In summer, the combination of strong eastward pulses in the current and upwelling at the capes favoured dispersal onto the midshelf of the Agulhas Bank. In winter, alongshore oscillations without the offshore displacement associated with upwelling. restricted offshore dispersal which caused surface particles to be retained inshore. Drifter trajectories show that both the eastward and westward nearshore current can link the inshore spawning grounds with the nursery grounds, offshore on the central Agulhas Bank; and that passive, neutrally buoyant material in the surface layer can reach the vicinity of the cold ridge in as little as eight days. The wind-driven processes of upwelling and coastal trapped waves (CTWs). and the influence of the greater shelf circulation are discussed as possible driving forces of variability in the currents off the Tsitsikamma coast. The occurrence of coastal trapped waves during thermal stratification appears to drive the jet-like, eastward pulses in the current. and results suggest that the propagation of CTWs may regulate and even enhance upwelling and downwelling along the Tsitsikamma coast.

It was previously thought that the South African chokka squid Loligo reynaudi is exclusively an inshore, shallow water spawner. Although spawning mostly within shallow bays (<60 m) the presence of squid eggs in trawls at depths up to 130 m indicates this species frequently makes use of deeper spawning areas on the mid-shelf. The extent of mid-shelf spawning (referred to as deep spawning) and the contribution to recruitment has yet to be assessed. Studies have shown mid-shelf bottom temperature to vary considerably from those inshore, suggesting chokka squid spawn in two very different oceanographic environments. Considering these apparent environmental differences, what leads to the mid-shelf environment becoming a suitable spawning habitat? Does a suitable benthic habitat, required for the attachment of egg pods, occur on the mid-shelf? These questions are not only important for determining the extent of deep spawning, but also to the understanding of factors “driving” deep spawning. The fate of deep spawned hatchlings is another unknown. It has been proposed that the main chokka squid inshore spawning grounds are positioned to exploit the net westward currents on the Eastern Agulhas Bank, i.e. paralarvae would be transported west from the hatching site to the cold ridge, an area of high primary and secondary productivity on the Central Agulhas Bank. This concept has come to be known as the Western Transport Hypothesis. Lagrangian ROMS-IBMs (regional ocean model system – individual-based model) predict the net westward transport of paralarvae from both the inshore and deep spawning grounds, to the cold ridge. These simulations were used to investigate the transport of hatchlings to the cold ridge feeding grounds before the exhaustion of yolk reserves. The fate of paralarvae on reaching the feeding grounds has not yet been investigated. This work has contributed new knowledge to our understanding of deep spawning and its role in recruitment. Specific aims of this study were to (1) determine the extent, range and importance of the deep spawning grounds relative to those inshore; (2) investigate the deep spawning ground habitat (Agulhas Bank mid-shelf) morphology and oceanographic environment; (3) determine the transport and survival of deep spawned hatchlings; and (4) investigate the origin and distribution of chokka squid paralarvae on the Agulhas Bank. The extent, depth range and importance of the deep spawning grounds, relative to those inshore was assessed using 23 years of demersal trawl survey data. Data for both the west and south coasts of South Africa were examined for egg capsules. No spawning was found on the west coast. Data showed that chokka squid preferred the Eastern Agulhas Bank for spawning. Spawning occurred not only inshore but also on the mid-shelf extending to depths of 270 m near the shelf edge. The majority of deep spawned eggs however, were found in the depth range 71-130 m. Squid egg density markedly decreased beyond 70 m, suggesting delineation between the inshore and deep spawning grounds. Total egg biomass calculations for depths shallower and deeper than 70 m indicated the coastal area to be strongly favoured, i.e. 82 vs. 18%. These results contest the commonly accepted notion that chokka squid is an inshore spawner and redefine the spawning grounds to extend across the shelf. Apart from an initial study investigating bottom temperature on the mid-shelf, very little is known about the deep spawning habitat. St Francis Bay, a commonly used spawning location, was chosen as a demonstration area for further study. The deep spawning grounds (71-130 m) were mapped and benthic habitat described from underwater video footage. A study investigating cross-shelf bottom conditions was undertaken off Thys Bay. CTD data were used to compare seasonal bottom temperature and oxygen on the St Francis Bay inshore and deep spawning grounds. Squid movement between the two spawning habitats was assessed using filament tagging. Predation and fishing pressure across the spawning grounds was reviewed. The mid-shelf benthic habitat was found to be similar to that inshore and available for spawning. Despite the generally colder bottom temperatures on the mid-shelf, this study showed that bottom temperature in deeper waters can at times be warmer than inshore. Although mid-shelf warming events lasted from a few hours to a number of days, they resulted in similar conditions to those on the inshore spawning grounds. It is likely these events act to expand or shift spawning habitat. The movement of squid between the two spawning habitats makes it possible for them to seek patches of warm bottom water with appropiate substrate. This suggests they are spawning habitat opportunists. Predation and fishing pressure appear to be higher on the inshore spawning grounds. It is feasible that this also forces spawners to seek out more favourable habitat offshore. An individual-based model was used to predict the fate of mid-shelf and inshore hatched paralarvae. Within the model, both the highly productive cold ridge and inshore spawning grounds were considered feeding or nursery areas. Paralarvae were released from six inshore and six deep spawning sites, spanning the coast between Port Alfred and Knysna. All paralarvae not reaching the feeding areas before the exhaustion of yolk-reserves (≤5 days), not retained within the feeding grounds (≥14 days), and not retained on the Agulhas Bank after exiting the feeding grounds were considered lost. This work illustrated the dependence of paralarval transport success on both spawning location and time of hatching, as established in earlier studies. The current IBM has expanded on initial work, emphasizing the importance of the cold ridge and inshore spawning grounds as nursery areas for deep and inshore spawned paralarvae, respectively. This work has highlighted the complex interactions between processes influencing recruitment variability for chokka squid. Possible relationships between periods of highest recruitment success and spawning peaks were identified for both spawning habitats. Based on the likely autumn increase in deep spawning off Tsitsikamma, and the beneficial currents during this period, it can be concluded deep spawning may at times contribute significantly to recruitment. This is particularly true for years where the cold ridge persists into winter. Data on chokka squid paralarval distribution are scarce. Paralarval distribution and abundance, in relation to Agulhas Bank oceanography, was investigated using bongo caught paralarvae and corresponding oceanographic data. Individual-based models (IBMs) were used to predict the origin or spawning site of the wild caught paralarvae, with reference to inshore versus deep spawning. Although failing to predict realistic points of origin, this study provided evidence to support a number of scenarios previously assumed to influence chokka squid recruitment. First is the possible influence of coastal upwelling on the retention, and hence spatial distribution, of paralarvae on the inshore spawning grounds. The second factor thought to impact recruitment is the loss of paralarvae from the Agulhas Bank ecosystem. This study confirmed the removal of paralarvae from the Eastern Agulhas Bank due to Agulhas Current boundary phenomena and resultant offshelf leakage. In addition, data suggested that the formation of the cold ridge could enhance retention on the Central Agulhas Bank, and so prevent offshelf leakage from the Central and Western Agulhas Bank. A synthesis of the main conclusions is presented. Implications of the findings and directions for future research are discussed.

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.

Σκοπός της παρούσας διατριβής, ήταν α) η μελέτη της συστηματικής των Κεφαλοπόδων και ειδικώτερα της κατανομής και των συναθροίσεών τους στο Θρακικό πέλαγος και τους κόλπους Στρυμωνικό, Συγγιτικό, Τορωναίο, Θερμαϊκό και β) η μελέτη του βιολογικού κύκλου των ειδών Loligo vulgaris (καλαμάρι), Illex coindetii (θράψαλο) και Sepietta oweniana. Το υλικό που χρησιμοποιήθηκε προήλθε κυρίως από 8 δειγματοληψίες (9/91 – 12/93) που πραγματοποιήθηκαν από το EΛ.ΚΕ.Θ.Ε., με τράτα βυθού, σε βάθη 17- 400 m. Για τη μελέτη της βιολογίας των 2 Τευθοειδών το υλικό συμπληρώθηκε με μηνιαία λήψη δειγμάτων (2/92 – 6/93) από την επαγγελματική αλιεία στο ΒΑ Αιγαίο. Συνολικά προσδιορίστηκαν 28 είδη Κεφαλοπόδων, από 8 διαφορετικές οικογένειες: Enoploteuthidae (1 είδος), Histioteuthidae (1 είδος), Loliginidae (3 είδη), Ommastrephidae (3 είδη), Sepiidae (3 είδη), Sepiolidae (9 είδη), Argonautidae (1 είδος) και Octopodidae (7 είδη). Πολυπαραγοντικές αναλύσεις των λογαριθμικά τροποποιημένων δεδομένων αφθονίας των ειδών ανά δειγματοληπτική σύρση, έδειξαν διαφοροποιήσεις στη δομή των συναθροίσεων των Κεφαλοπόδων κατά κύριο λόγο με το βάθος και σε μικρότερο βαθμό με την εποχή και τη γεωγραφική περιοχή δειγματοληψίας. Στα είδη I. coindetii και S. oweniana διαπιστώθηκε αναπαραγωγική δραστηριότητα καθόλη τη διάρκεια του χρόνου, με μέγιστη ένταση αντίστοιχα το φθινόπωρο και χειμώνα – άνοιξη. Η αναπαραγωγική περίοδος για το νηριτικό L. vulgaris ήταν σχετικά πιό περιορισμένη (μέσα χειμώνα - αρχές φθινοπώρου) με μέγιστη ένταση την άνοιξη. Η ηλικία των 2 τευθοειδών, που εκτιμήθηκε απο την ανάγνωση των αυξητικών δακτυλίων σε στατολίθους, δεν ξεπερνά τους 14 μήνες ενώ ο ημερήσιος ρυθμός αύξησης του μανδύα φτάνει για το καλαμάρι τα 2-2,5 mm και για το θράψαλο τα 0,5-0,6 mm. Οι Οστειχθείς και τα Κεφαλόποδα αποτελούσαν τις πιο συχνά εμφανιζόμενες λείες στο στομαχικό περιεχόμενο των δύο Τευθοειδών, ενώ τα Καρκινοειδή την προτιμώμενη λεία για το είδος S. oweniana. / The twofold aim of this study was: firstly to contribute to the knowledge of the cephalopod species taxonomy in the Greek Seas, focusing particularly on faunistic composition at the N. Aegean (ΝΕ Mediterranean), and secondly to investigate the life history patterns of the species Loligo vulgaris, Illex coindetii and Sepietta oweniana. Faunistic study was based on samples collected from the N. Aegean Sea (N>39ο 50΄) during eight trawl surveys (9/91 – 12/93), carried out at depths 17 - 400 m. For the study of the two squid species biology, additional monthly samples were collected from commercial fishery in NE Aegean Sea during the period February 1992 - June 1993. In all, 28 species of cephalopods belonging to 8 families were identified including Enoploteuthidae (1 species), Histioteuthidae (1 species), Loliginidae (3 species), Ommastrephidae (3 species), Sepiidae (3 species), Sepiolidae (9 species), Argonautidae (1 species) and Octopodidae (7 species). To detect zonation patterns in cephalopod community structure, multivariate analyses of species abundance data per haul were performed. Considerable variability was shown in assemblage structure, determined primarily by depth, and to a lesser extent, by geographical location and season. Intermittent terminal spawning pattern has been shown for all 3 examined species. Spawning occurs throughout the year for I. coindetii and S. oweniana in the N. Aegean Sea, peaking respectively in autumn and winter-spring, as indicated by minimum ML50 values. The breeding season of neritic L. vulgaris extends from late winter to early autumn, with spawning intensity varying among years, due to variation in temperature and population age structure. Age of both teuthoid species, estimated by growth increment counts on statoliths, did not exceed 14 months. Daily growth rate (DGR) reached 2-2,5 mm in L. vulgaris and 0,5-0,6 mm in I. coindetii. L. vulgaris and I. coindetii feed primarily on fishes and cephalopods in the N. Aegean Sea, whereas S. oweniana on crustacean and fishes.