Erratic and highly variable catches in the South African chokka squid Loligo vulgaris reynaudii fishery, cause socio-economic hardship for the industry and uncertainty for resource managers. Catch forecasting can reduce this problem as it is believed that catch variability is strongly influenced by environmental factors. In this study, data were collected at varying temporal and spatial scales. Data for the hourly time-scale study were collected from 1996-1998, aboard commercial vessels, whilst for the longer time-scales, data were extracted for Kromme Bay (a single fishing area) from existing databases (1991-1998) that were comprised of compulsory catch returns and oceanographic data. The environment-catch relationship for chokka squid on the inshore spawning grounds was then investigated using multiple correlation and regression analysis, analysis of variance, contingency table analysis and cross-correlation statistical techniques. This simple, direct, 'black box' statistical approach was relatively successful in developing a predictive capability. On a short time-scale (hourly), the regression model accounted for 32% of the variability in catch, with turbidity the main determinant (13%). On a daily monthly time-scale, the best prediction model was on a monthly scale, accounting for 40% of the variability in catch. The principal determinant, bottom temperature anomaly (11 %), was found to lag one month forward. Seasonal and diel catch variations induced changes in the relative importance of turbidity, water temperature and wind direction on catches. A strong, positive relationship was found between easterly winds (which cause upwelling) and catch, particularly in summer. Catch rates, however, decreased with an increase in turbidity. The correlation between temperature and catch was generally negative, however, higher catches were associated with a temperature range of 13-18°C. Highest catch rates were associated with easterly winds, zero turbidity conditions and sea surface temperatures from 15.0-16.9°C. Selected case studies (in situ observations) suggested that upwelling and turbidity events act as environmental triggers for the initiation or termination of the spawning process, respectively. A holistic approach is required to improve predictive capability of chokka squid abundance. Although short-term predictability remains essential (i.e. hourly-scale), future research should concentrate on long-term prediction models (e.g., monthly time-scales) involving greater spatial variation, which are the most important for management.

Several studies suggest the environment influences chokka squid catches which are mostly based on the successful formation of inshore spawning aggregations. None of the evidence, however, is direct observation. Acoustic telemetry offers a means to determine the response of spawners to changes in the environment and insight into the behaviour of spawning squid. A hexagonal array of VR2 receivers deployed 500 m apart was deemed to be ideal to monitor the movement patterns of squid on the spawning sites. In isothermic conditions, an area up to 1.28 km2 could be monitored as there was an approximate 50 m overlap in individual VR2 receiver range. In thermocline conditions however, “acoustic dead zones” as wide as 350 m may have existed between VR2 receivers, limiting the performance of this configuration. Similarly benthic turbidity events would also decrease detection range and limit performance. A hexagonal array of VR2 receivers was moored in Kromme Bay on and around active spawning aggregations during the squid fishery closed seasons of November 2003, 2004, 2005 and 2006. Squid were caught on jigs and tagged with V9 acoustic pressure telemetry transmitters. A total of 45 animals were tagged. Presence-absence analysis identified three general behaviours: (1) arrival at dawn and departure after dusk, (2) a continuous and uninterrupted presence for a number of days and (3) presence interrupted by frequent but short periods of absence. Overall, the data suggests frequent migrations between spawning aggregations and offshore feeding grounds. The pressure sensor data showed both males and females stayed persistently near the seabed during the day, but at night, this pattern was broken with common activity higher up in the water column. The squid did not remain exclusively in the water column and regularly made excursions to the seabed. CTD and temperature data indicated the intrusion of a cold bottom layer due to upwelling at the monitored spawning sites on a number of occasions. The formation of spawning aggregations appears to be triggered by upwelling events and spawning behaviour, once initiated, disrupted by upwelling events with a rapid onset, possibly due to an inability to adapt physiologically over such a short time period.

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.