Life-history parameters of adult females of Merluccius capensis (Gadidae) off the south coast of South Africa.

This study explores the life-history parameters of female Merluccius capensis off South Africa (N = 1819) during 2014-2016, including gonadosomatic index (GSI), length-at-maturity, length-weight relationships, and condition indices (relative condition [k] and Fulton's condition factor [K]). We detected weak indications of two peaks of spawning within the year, the first in austral autumn from March to May, whereas the other in austral spring around August. GSI was slightly higher in spring and autumn, though still low at all maturity stages (≤7%), though the opposite was true for the actively spawning stage (≥7%) as well as access to less such data during winter- and summertime. The length (L) at 50% maturity was around 38 cm (L50), though differences occurred between the two applied staging methods, histology and visual (macroscopic) classification, when L approached infinity. The latter method presented underestimated length at maturity values at the 75 and 95 percentiles (48 and 60 cm) compared to the corresponding percentiles given by histology (50 and 65 cm). There were trivial across-method differences in L50. However, we found a clear reduction in L50 in view of published information in prior years when this estimate was 48 (1985), 42 (2008), 53 (2011), and 24.8 (2015) cm. Overall, L explained 90% of the variation in whole body weight (W). As the bootstrapped, grand mean growth coefficient was b = 2.98, indicating a slight allometric growth function, there were no significant variations between years, though an isometric growth existed for 2016 with b = 3.0, whereas for 2014 and 2015 this b was 2.98 and 2.93, respectively. In terms of demography, females <60 cm generally showed isometric growth (b = 3) as opposed to allometric growth (b = 2.95) at >60 cm. The relative condition index (k = 1) exhibited higher values than Fulton's K, which was 0.80. Overall, the maternal stock of M. capensis along the south coast seems to be in good condition and likely spawns throughout the year, but we found that the macroscopic data tend to give biased maturity ogives.

Spatio-temporal genetic structure and the effects of long-term fishing in two partially sympatric offshore demersal fishes.

Environmental gradients have been shown to disrupt gene flow in marine species, yet their influence in structuring populations at depth remains poorly understood. The Cape hakes (Merluccius paradoxus and M. capensis) are demersal species co-occurring in the Benguela Current system, where decades of intense fishing resulted in severely depleted stocks in the past. Previous studies identified conflicting mtDNA genetic substructuring patterns and thus contrasting evolutionary trajectories for both species. Using 10 microsatellite loci, the control region of mtDNA and employing a seascape genetics approach, we investigated genetic connectivity and the impact of prolonged exploitation in the two species, which are characterized by different patterns of fishing pressure. Three consecutive years were sampled covering the entire distribution (N = 2100 fishes). Despite large estimated population sizes, both species exhibited low levels of contemporary genetic diversity (0.581 < HE  < 0.692), implying that fishing has had a significant impact on their genetic composition and evolutionary trajectories. Further, for M. paradoxus, significant temporal, but not spatial, divergence points to the presence of genetic chaotic patchiness. In contrast, M. capensis exhibited a clear latitudinal cline in genetic differentiation between Namibia and South Africa (FST  = 0.063, P < 0.05), with low (0.2% per generation) estimates of contemporary gene flow. Seascape analyses reveal an association with bathymetry and upwelling events, suggesting that adaptation to local environmental conditions may drive genetic differentiation in M. capensis. Importantly, our results highlight the need for temporal sampling in disentangling the complex factors that impact population divergence in marine fishes.

Environmental effects on cephalopod population dynamics: implications for management of fisheries.

Cephalopods are a relatively small class of molluscs (~800 species), but they support some large industrial scale fisheries and numerous small-scale, local, artisanal fisheries. For several decades, landings of cephalopods globally have grown against a background of total finfish landings levelling off and then declining. There is now evidence that in recent years, growth in cephalopod landings has declined. The commercially exploited cephalopod species are fast-growing, short-lived ecological opportunists. Annual variability in abundance is strongly influenced by environmental variability, but the underlying causes of the links between environment and population dynamics are poorly understood. Stock assessment models have recently been developed that incorporate environmental processes that drive variability in recruitment, distribution and migration patterns. These models can be expected to improve as more, and better, data are obtained on environmental effects and as techniques for stock identification improve. A key element of future progress will be improved understanding of trophic dynamics at all phases in the cephalopod life cycle. In the meantime, there is no routine stock assessment in many targeted fisheries or in the numerous by-catch fisheries for cephalopods. There is a particular need for a precautionary approach in these cases. Assessment in many fisheries is complicated because cephalopods are ecological opportunists and stocks appear to have benefited from the reduction of key predator by overexploitation. Because of the complexities involved, ecosystem-based fisheries management integrating social, economic and ecological considerations is desirable for cephalopod fisheries. An ecological approach to management is routine in many fisheries, but to be effective, good scientific understanding of the relationships between the environment, trophic dynamics and population dynamics is essential. Fisheries and the ecosystems they depend on can only be managed by regulating the activities of the fishing industry, and this requires understanding the dynamics of the stocks they exploit.

Management and conservation implications of cryptic population substructure for two commercially exploited fishes (Merluccius spp.) in southern Africa.

Genomic information can aid in the establishment of sustainable management plans for commercially exploited marine fishes, aiding in the long-term conservation of these resources. The southern African hakes (Merluccius capensis and M. paradoxus) are commercially valuable demersal fishes with similar distribution ranges but exhibiting contrasting life histories. Using a comparative framework based on Pool-Seq genome-wide SNP data, we investigated whether the evolutionary processes that shaped extant patterns of diversity and divergence are shared among these two congeneric fishes, or unique to each one. Our findings revealed that M. capensis and M. paradoxus show similar levels of genome-wide diversity, despite different census sizes and life-history features. In addition, M. capensis shows three highly structured geographic populations across the Benguela Current region (one in the northern Benguela and two in the southern Benguela), with no consistent genome-environment associations detected. In contrast, although population structure and outlier analyses suggested panmixia for M. paradoxus, reconstruction of its demographic history suggested the presence of an Atlantic-Indian Ocean subtle substructuring pattern. Therefore, it appears that M. paradoxus might be composed by two highly connected populations, one in the Atlantic and one in the southwest Indian Ocean. The reported similar low levels of genomic diversity, as well as newly discovered genetically distinct populations in both hake species can thus assist in informing and improving conservation and management plans for the commercially important southern African Merluccius.

Remarkably low mtDNA control region diversity in an abundant demersal fish.

Cape hake, Merluccius paradoxus, is a valuable commercially exploited demersal species. Using the 5' mtDNA control region we show that 96% of 1013 fishes sampled over a three-year period share one of two dominant haplotypes; 19 haplotypes were recovered in total, suggesting a genetically homogenous population of fish. Accordingly, haplotype and nucleotide diversities are low (h = 0.53, pi = 0.0014); an asymptotic haplotype accumulation curve suggests that few additional haplotypes exist. Comparing h and pi with other fish species shows that M. paradoxus and other southern African fish species have remarkably low genetic diversity values compared with other global marine fishes. Despite low genetic variability, frequency differences among M. paradoxus haplotypes suggest weakly structured populations between Namibia and South Africa. However, given the remarkably homogeneous mtDNA population genetic structure between fishes sampled along 1800 km, it is clear that faster evolving markers such a microsatellites are also needed before inferences can be made regarding stock identification and management of this species.

Sperm storage and mating in the deep-sea squid Taningia danae Joubin, 1931 (Oegopsida: Octopoteuthidae).

Spermatangium implantation is reported in the large oceanic squid Taningia danae, based on ten mated females from the stomachs of sperm whales. Implanted spermatangia were located in the mantle, head and neck (on both sides) or above the nuchal cartilage, under the neck collar and were often associated with incisions. These cuts ranged from 30 to 65 mm in length and were probably made by males, using the beak or arm hooks. This is the first time wounds facilitating spermatangium storage have been observed in the internal muscle layers (rather than external, as observed in some other species of squid). The implications of these observations for the mating behavior of the rarely encountered squid T. danae are discussed.

Mitochondrial DNA analyses of the Cape hakes reveal an expanding, panmictic population for Merluccius capensis and population structuring for mature fish in Merluccius paradoxus.

The Cape hake species, Merluccius capensis and Merluccius paradoxus are the most important resource of the South African and Namibian demersal fishery, but it is unclear whether there is a single population of each shared by both countries. We analysed the population structure and evolutionary history of these two species using the variable 5' region of the mtDNA control region for 311 specimens of M. capensis and 333 specimens of M. paradoxus sampled between Lüderitz (southern Namibia) to south of Cape Point (South Africa). 107 haplotypes for M. capensis and eight haplotypes for M. paradoxus were recovered. AMOVA and pairwise Phi(st) analyses revealed no structure in M. capensis, however significant genetic differentiation between Namibian and South African 'populations' was detected for M. paradoxus. This was only restricted to mature fish older than 3 and 4 years and not for juvenile fish younger than 3 years. Analyses reveal that M. capensis has undergone population expansion (Fu's Fs=-26.65, P<0.001), possibly within the last 4500-23,000 years, whereas M. paradoxus has not. Our study highlights the utility of genetic markers to unravel the evolutionary history of sympatric species, as well as addressing management issues within regions where commercially valuable fish stocks are shared between nations.