Distribution,body size and biology of the megamouth shark Megachasma pelagios

We compiled historical reports of megamouth sharks Megachasma pelagios (mostly fishery by-catch and strandings) from 1976 to 2018 (n = 117) and found that they are distributed globally (highest latitude, 36°) with three hotspots: Japan, Taiwan and the Philippines. Despite possible biases due to variability in fishing effort, more individuals were reported at higher latitudes in the summer, suggesting seasonal, latitudinal migrations. Sex ratios were female-biased in Japan, but more even in Taiwan and the Philippines, suggesting some sexual segregation. Females (total length, L_T = 3.41–7.10 m) were larger than males (L_T = 1.77–5.39 m) and matured at a larger L_T (5.17 m) than males (4.26 m). Also, we reviewed the systematics, feeding ecology and swimming behaviour of Megachasma pelagios based on the literature. Our review shows that, compared with their morphology, anatomy and genetics, behavioural ecology of this species remains largely unknown and electronic tagging studies are warranted.     

Catch composition,reproductive biology and diet of the bramble shark Echinorhinus brucus (Squaliformes: Echinorhinidae) from the south‐eastern Arabian Sea

Fishery and biological data are presented for the poorly known bramble shark Echinorhinus brucus (Squaliformes: Echinorhinidae), from the deep waters of the south‐eastern Arabian Sea. A total of 5318 individuals from by‐catch landings of deep‐water bottom set longlines, gillnets and shrimp trawl fisheries operating at depths of 200–1200 m were recorded between January 2008 and December 2011 at the Kochi Fisheries Harbour (Kerala). A total of 431 individuals, from 46 to 318 cm total length (L_T) and 0·8 to 132 kg total mass (M_T), were examined to determine biological data for this species. The L_T at which 50% were mature (L_T50) for females and males was estimated at 189 and 187 cm L_T. Litter size ranged from 10 to 36 and size at birth was between 42 and 46 cm L_T. Dietary analysis of stomach contents revealed E. brucus feeds on a variety of prey including crustaceans (69% index of relative importance, I_RI), teleosts (25·8% I_RI), cephalopods (1·7% I_RI) and elasmobranchs (0·7% I_RI). This study provides the first detailed biological data for this species and also highlights the extent of the by‐catch fishery for this species in Indian waters.     

The diet of Clinitrachus argentatus (Blennioidei: Clinidae) in the northern Aegean and Marmara Seas

Differences in the diet composition of small (28–42 mm total length, L_T) and large (43–75 mm L_T) individuals of the cline Clinitrachus argentatus (Clinidae) were evident. Smaller individuals preyed mainly on copepods, whereas larger individuals ate mainly amphipods, isopods, brachyurans and polychaetes.     

Biological observations on the broadfin shark Lamiopsis temminckii (Carcharhiniformes: Carcharhinidae)

Biological information was collected from 214 individuals of the broadfin shark Lamiopsis temminckii measuring 418 to 1782 mm total length, L_T. Size at maturity (L₅₀) for females and males was estimated at 1430 and 1368 mm L_T, respectively, while mature and gravid females were observed from 1350 mm L_T with litter sizes 2–8 and size at birth 418–650 mm L_T. Analysis of stomach contents revealed a variety of prey, primarily crustaceans (54·0%), teleosts (42·7%) and cephalopods.     

Reproductive biology of the fat snook Centropomus parallelus Poey, 1860 (Teleostei,Centropomidae) and implications for its management in the southern Atlantic Ocean

The reproductive biology of Centropomus parallelus was described from 589 individuals captured in estuarine and coastal waters in Southern Brazil. Length-frequency distribution showed the dominance of males in smaller length-classes (132–290 mm L_T), whereas females were dominant in larger length-classes (>290 mm L_T). Total length at maturity (L₅₀) was 180 mm L_T and corresponded to 29% of the maximum length recorded. Histological sections revealed one hermaphrodite (205 mm L_T) and few immature females. Life history traits provided herein can contribute to sustainable fisheries management practices.     

Habitat use and early winter movements by juvenile Atlantic cod in a coastal area of Newfoundland

Acoustic telemetry was used to monitor the movements and landscape scale habitat use of age 2–3 year juvenile Atlantic cod Gadus morhua in Newman Sound, a coastal fjord of Newfoundland, during late autumn and early winter (24 October 1999 to 22 January 2000). Substratum, bathymetric relief and depth use were studied to determine if it differed from the pattern expected given an absence of selectivity (habitat use proportional to habitat availability). Prior to winter migrations, most Atlantic cod maintained small home ranges (0·5–33·4 ha, median = 2·1 ha) although a few individuals moved more widely. No relationship between total length (L_T) and home range size was detected. In Inner Newman Sound, Atlantic cod occupied depths of 10–29 m more than expected given availability, while depths of 0–9 and 40–59 m were underutilized. No significant relationship between depth and L_T was detected. Areas of medium (5–10%) or high (>10%) bathymetric relief and boulder or kelp habitats were used significantly more than expected given the availability of these habitats. Sand and eelgrass substrata were underutilized given availability, although many Atlantic cod used sandy‐bottomed areas to some degree. Flexibility in habitat use by the individuals that were studied suggested reduced predation risk relative to younger conspecifics. Winter migrations to deeper water beyond Newman Sound began in mid‐November, coinciding with the disappearance of the thermocline, and continued until 27 December. Approximately 30% of monitored individuals did not migrate and maintained their home ranges into the winter season. No significant differences in L_T between migrating and resident groups were detected, however, the condition of migrating fish was significantly higher than in resident fish. This finding supports the theory that feeding history plays a role in the decision to undertake migration.     

Interaction between migration behaviour and estuarine mortality in cultivated Atlantic salmon Salmo salar smolts

Migration behaviour and estuarine mortality of cultivated Atlantic salmon Salmo salar smolts in a 16 km long estuary were studied using two methods: (1) acoustic telemetry and (2) group tagging in combination with trap nets. Progression rates of surviving individuals through the estuary were relatively slow using both methods [0·38 L_T (total length) s^?1 v. 0·25 L_T s^?1]. In 2012, the progression rate was slow from the river to the estuary (0·55 L_T s^?1) and the first part of the estuary (0·31 L_T s^?1), but increased thereafter (1·45–2·21 L_T s^?1). In 2013, the progression rate was fast from the river to the estuary (4·31 L_T s^?1) but was slower thereafter (0·18–0·91 L_T s^?1). Survival to the fjord was higher in 2012 (47%) compared to 2013 (6%). Fast moving individuals were more likely to migrate successfully through the estuary compared to slower moving individuals. Adult recapture of coded‐wire‐tagged S. salar was generally low (0·00–0·04%). Mortality hot spots were related to topographically distinct areas such as the river outlet (in 2012) or the sill separating the estuary and the fjord (in 2013). At the sill, an aggregation of cod Gadus morhua predating on cultivated smolts was identified. The results indicate that slow progression rates through the estuary decreases the likelihood of smolts being detected outside the estuary. The highly stochastic and site‐specific mortality patterns observed in this study highlight the complexity in extrapolating mortality patterns of single release groups to the entire smolt run of wild S. salar.     

Skeletal ontogeny of the vertebral column and of the fins in shi drum Umbrina cirrosa (Teleostei: Perciformes: Sciaenidae), a new candidate species for aquaculture

The osteological development of the vertebral column and fins in shi drum Umbrina cirrosa was studied in order to improve knowledge for its introduction in Mediterranean aquaculture. The osteological development was studied in 171 individuals, of total length (L_T) from 2·7 to 30·2 mm that were reared under the mesocosm technique. Vertebral ontogeny starts at 3·4 and 4·0 mm L_T, with the formation of the first cartilaginous neural and haemal arches, and spines, respectively, and is completed with the full attainment of epicentrals (12·5 mm L_T). The formation of vertebral centra occurs between 4·1 and 7·4 mm L_T. Pectoral supports are the first fin elements to develop (3·0 mm L_T), followed by those of the caudal fin (3·8 mm L_T), pelvic fin (3·9 mm L_T) and finally by those of the dorsal and anal fins (4·5 mm L_T). The caudal fin is the first to develop fin rays and attain the full count of principal fin rays (4·5–6·8 mm L_T), but the last to be fully completed with the formation of procurrent fin rays (6·9–17·5 mm L_T). The next fins starting to present rays are the dorsal (5·3 mm L_T) and the pectoral fins (5·6 mm L_T), while the anal and pelvic fins are the last (5·7 mm L_T). Following the caudal principal fin rays (6·8 mm L_T), the dorsal, anal (6·9 mm L_T), pelvic (7·4 mm L_T) and pectoral fins (9·8 mm L_T) are the next with fully completed ray counts. Aggregation of qualitative changes, such as the appearance of cartilages, the beginning and the complement of the ossification process and the full complement of elements in U. cirrosa were measured as cumulative frequency counts. These measurements reveal three ontogenetic intervals: one very developmentally active period during early life stages (from 3 to 5·9 mm L_T), a second slower developmental period (from 6·0 to 8·9 mm L_T) and finally a period of ontogeny more focused on structure refinement up to metamorphosis and settlement (>9·0 mm L_T).     

Diel vertical migrations of bathypelagic perch fry

The behaviour of young‐of‐the‐year (YOY) perch Perca fluviatilis as a dominant species in the assemblage of fry in the pelagic of Slapy Reservoir (Czech Republic), was studied during late May and mid‐June 2002 using acoustic methods and complementary net catches. During the day, perch fry were present simultaneously in littoral, epipelagic and bathypelagic habitats. Bathypelagic perch fry, forming a scattering layer, migrated vertically each day between the epilimnion and hypolimnion, with an amplitude of 11·0 m in May and 12·5 m in June. At dusk, the migratory bathypelagic fry mixed in the epilimnion with non‐migrating epipelagic fry and spent the night close to the thermocline (abundance maximum at 3–4 m in May, 0–2 m in June). In June, shoaling behaviour by some of the bathypelagic perch fry was also observed: the shoaling fry remained higher in the water column than the non‐shoaling fry. Both depths of the scattering layer and the depths of the fry shoals were strongly controlled by the light intensity. The contribution of the bathypelagic part of the population to the total numbers of pelagic perch fry decreased from 28·1% in May to 4·7% in June, while the density of all pelagic perch fry increased (c. 96 000 individuals ha^?1 in May and 142 000 individuals ha^?1 in June). In May, the bathypelagic (average total length, L_T, 11·9 mm) and epipelagic (average L_T 14·6 mm) perch fry differed in size while, in June, the epipelagic fry were divided into two distinct size groups. The more abundant group, of small epipelagic perch fry (average L_T 14·6 mm), was similar in size to the bathypelagic fry (average L_T 14·6 mm) while the less abundant group, of larger epipelagic fry (average L_T 34·4 mm), was similar in size to littoral perch fry (average L_T 35·0 mm). The results suggest that in perch fry three different survival strategies with different risks can be used in the same locality, time and year.     

Distribution of Sprattus sprattus phalericus (Risso, 1827) and zooplankton near the Black Sea redoxcline

Understanding what environmental drivers influence marine predator–prey relationships can be key to managing and protecting ecosystems, especially in the face of future climate change risks. This is especially important in environments such as the Black Sea, where strong biogeochemical gradients can drive marine habitat partitioning and ecological interactions. We used underwater video recordings in the north-eastern Black Sea in November 2013 to observe the distribution and behaviour of the Black Sea sprat (Sprattus sprattus phalericus, Risso 1827) and its zooplankton prey. Video recordings have shown that the Black Sea sprat S. sprattus phalericus tolerates severely hypoxic waters near the redoxcline. The school was distributed in the 33–96 m layer [oxygen concentration (O₂) 277–84 μmol L⁻¹]. Some individuals were observed to leave the school and descended 20 m deeper for foraging on copepods in the 119–123 m layer (O₂ 12–10 μmol L⁻¹). Zooplankton appeared concentrated on the upper boundary of the suboxic zone (O₂ < 10 μmol L⁻¹). No zooplankton were observed below O₂ 6–7 μmol L⁻¹ (128 m). Understanding the ability of this species to tolerate low oxygen waters is crucial to predicting future responses to natural and anthropogenic changes in hypoxia.     

Population structure and residency of whale sharks Rhincodon typus at Utila,Bay Islands,Honduras

There were 479 reported whale shark Rhincodon typus encounters between 1999 and 2011 at the island of Utila, which forms part of the Meso‐American Barrier Reef System (MBRS) in the western Caribbean Sea. The majority of R. typus were found to feed on small bait fish associated with various tuna species. Ninety‐five individual R. typus, ranging from 2 to 11 m total length (L_T), were identified through their unique spot patterns. A significant male bias (65%) was present. There was no significant difference between the mean ± s.d . L_T of female (6·66 ± 1·65 m) and male (6·25 ± 1·60 m) R. typus. Most R. typus were transient to Utila, with 78% sighted only within a single calendar year, although some individuals were sighted in up to 5 years. Mean residency time was modelled to be 11·76 days using maximum likelihood methods.     

Acoustic ontogeny of a teleost

The influence of growth on the intensity and variability of acoustic echoes from individuals and groups of a teleosts was quantified using Donaldson trout (rainbow–steelhead hybrid) Oncorhynchus mykiss. Fish growth was linear in total length (L_T) and quadratic in mass. Dorsal swimbladder area increased exponentially with L_T. Allometric growth ratio (i.e. k) values of swimbladder length linearly increased with L_T. Average swimbladder volumes occupied 3–6% of fish body volume and increased exponentially with L_T. The aspect angle that resulted in the maximum average acoustic intensity from the group shifted from 80 to 86° through the experimental period. Mean echo intensities increased at both 38 and 120 kHz as mean L_T increased. Predicted echo intensities at 38 kHz exceeded that at 120 kHz at L_T <150 mm but were less than that predicted at 120 kHz at L_T >280 mm. Generalized additive mixed models using L_T, swimbladder angles and lateral elongation ratios of fish bodies were better predictors of echo intensities than L_T alone.     

Reproductive biology of the greeneye spurdog Squalus chloroculus (Squaliformes,Squalidae)

The reproduction of the greeneye spurdog Squalus chloroculus was studied based on animals caught in the multispecies and multi‐gear southern and eastern scalefish and shark fishery on the upper continental slope off southern Australia. One hundred and ninety‐nine females (502–990 mm, total length, L_T) and 189 males (515–810 mm L_T) were examined. The female reproductive cycle, based on 41 breeding animals, is continuous and triennial, with the pregnancy period estimated to be 31–34 months, seasonal and synchronous with the ovarian cycle; a third of the breeding female population is estimated to give birth between September and December each year. The estimated L_T at which 50% of females are mature is 799 mm (95% c.i. : 794, 804), whereas the L_T at which 50% are maternal is 825 mm (95% c.i. : 817–833), but these estimates are probably biased by the phenomenon of apparent change of L_T at maternity and L_T at maturity following severe length‐selective fishing mortality. Litters ranged from four to 15 embryos with a 1:1 sex ratio, and litter size increased with maternal length. The breeding cycle of males is neither seasonal nor synchronous with the female cycle. The estimated L_T of males where 50% are mature was 629 mm (95% c.i. : 603, 645).     

Biological observations on the bristly catshark Bythaelurus hispidus from deep waters off the south‐west coast of India

Biological data are presented for the poorly known bristly catshark Bythaelurus hispidus based on specimens collected from the by‐catch of the commercial deep‐sea shrimp trawl fishery operating in the Arabian Sea at depths of 200–500 m off the south‐west coast of India. One hundred and sixty‐two individuals, which ranged from 120 to 366 mm total length (L_T), were collected for this study. Size‐at‐maturity (L₅₀) for females and males was estimated at 252 and 235 mm L_T, respectively. The reproductive mode of B. hispidus was aplacental viviparity, which is the rarest reproductive mode within the Scyliorhinidae and is considered to be the most advanced of the three reproductive modes occurring within this family. Dietary analysis of stomach contents revealed B. hispidus feeds on a variety of prey, primarily fishes.     

Temporal and latitudinal comparisons of reproductive parameters in a heavily exploited shark,the bonnethead,Sphyrna tiburo (L. 1758), in the southern Gulf of Mexico

In the southern Gulf of Mexico, the bonnethead shark, Sphyrna tiburo, is one of the most frequently captured species in landings of small-scale fisheries. Based on the analysis of two fishery-dependent sampling periods (1993–1994 and 2007–2014), this study aimed to determine reproductive parameters and identify temporal differences between the two time periods. In the first sampling period, 776 males and 352 females with a size range of 28.0–120.0 cm total stretched length (L_T) were analysed, and in the second sampling period, 387 males and 432 females with a size range of 28.0–122.0 cm L_T were analysed. The size at 50% maturity in the second sampling period was significantly different between sexes, 82.6 cm L_T for females and 73.8 cm L_T for males (no estimation was possible for the first sampling period). The size at 50% maternity was not different between sampling periods, 97.3 cm L_T for the first sampling period and 99.0 cm L_T for the second sampling period. Litter size varied from 3 to 19 embryos and the average was not statistically different in both periods, 10.1 (S.D. = 3.8) for the first sampling period and 11.3 (S.D. = 3.5) for the second sampling period. The female reproductive cycle is asynchronous, and it seems to be annual, with a gestation period of 5–6 months, and a consecutive ovarian cycle and gestation period. Temporal (between sampling periods) and latitudinal (southern Gulf versus northern regions) variations occur in the synchronicity of the reproductive cycle, temporal variation in the relationship between maternal length and litter size, and latitudinal variation in average size of mature sharks.     

New records of angular rough sharks Oxynotus centrina in the coastal waters of Malta,with observations on post‐capture resilience and release behaviour

One male and one female angular rough sharks Oxynotus centrina were caught south‐east of Malta between May and June at a depth of 60–100 m < 5 km from shore. The immature female (total length, L_T 565 mm) was landed dead but the male (535 mm L_T) was found alive. This communication presents important biological observations on post‐capture recovery and release behaviour of this species.     

Incident radiation and the allocation of nitrogen within Arctic plant canopies: implications for predicting gross primary productivity

Arctic vegetation is characterized by high spatial variability in plant functional type (PFT) composition and gross primary productivity (P). Despite this variability, the two main drivers of P in sub‐Arctic tundra are leaf area index (L_T) and total foliar nitrogen (N_T). L_T and N_T have been shown to be tightly coupled across PFTs in sub‐Arctic tundra vegetation, which simplifies up‐scaling by allowing quantification of the main drivers of P from remotely sensed L_T. Our objective was to test the L_T–N_T relationship across multiple Arctic latitudes and to assess L_T as a predictor of P for the pan‐Arctic. Including PFT‐specific parameters in models of L_T–N_T coupling provided only incremental improvements in model fit, but significant improvements were gained from including site‐specific parameters. The degree of curvature in the L_T–N_T relationship, controlled by a fitted canopy nitrogen extinction co‐efficient, was negatively related to average levels of diffuse radiation at a site. This is consistent with theoretical predictions of more uniform vertical canopy N distributions under diffuse light conditions. Higher latitude sites had higher average leaf N content by mass (N_M), and we show for the first time that L_T–N_T coupling is achieved across latitudes via canopy‐scale trade‐offs between N_M and leaf mass per unit leaf area (L_M). Site‐specific parameters provided small but significant improvements in models of P based on L_T and moss cover. Our results suggest that differences in L_T–N_T coupling between sites could be used to improve pan‐Arctic models of P and we provide unique evidence that prevailing radiation conditions can significantly affect N allocation over regional scales.     

Reproductive biology of the southern thorny skate Amblyraja doellojuradoi (Chondrichthyes,Rajidae)

The total lengths (L_T) of 193 males (209–556 mm) and 130 females (275–515 mm) of Amblyraja doellojuradoi, a commercial by‐catch species on the Argentinean continental shelf, which are increasingly retained, were analysed. No sexual dimorphism was observed in the L_T at which 50% of individuals were sexually mature; males matured at 448 mm and females at 411 mm, c. 80 and 82% of maximum L_T. The hepato‐somatic index was similar among sexes, but significantly different between maturity stages, being lower in mature than immature specimens. Males had no seasonal difference in the hepato‐somatic index and females had the lowest index in autumn. The gonado‐somatic index was lower in males than in females and significantly higher in mature than immature specimens of both sexes. Males had the highest index in autumn and females had no seasonal difference. Collectively, these results would indicate that A. doellojuradoi breeds in autumn.     

Variation in the diet of the Patagonian toothfish with size, depth and season around the Falkland Islands

The ontogenetic and seasonal variations in the feeding spectrum were studied in 756 specimens of the Patagonian toothfish Dissostichus eleginoides (16–159 cm total length, L_T) collected on the shelf, continental slope and bathyal waters (67–1960 m, depth range) around the Falkland Islands between April 1999 and August 2002. On the shelf, small toothfish (<40 cm L_T) were active predators taking mostly one relatively large prey item at a time (mainly near‐bottom Patagonotothen ramsayi and Loligo gahi). Medium‐size toothfish (40–60 cm L_T) fed on the same prey, but the number of prey items increased to 1–2 items per fish. Large toothfish (>60 cm L_T) switched their diet to other large pelagic fishes occurring near the bottom (Macruronus magellanicus and Micromesistius australis australis), again taking mostly one prey item at a time. The diet of medium‐size D. eleginoides on the shelf varied seasonally depending on the abundance and migrations of the major prey species. Patagonotothen ramsayi was abundant in the diet throughout the year, whereas L. gahi appeared only from February to October during its offshore seasonal migrations to the depth range of D. eleginoides. During November to January, L. gahi migrated inshore to spawn and disappeared from the toothfish diet, being substituted by M. australis australis which dispersed on the shelf after spawning. After its ontogenetic descent to the lower part of the continental slope (500–1000 m depths), toothfish took less active (than on the shelf) fishes such as Antimora rostrata whilst also feeding on active near‐bottom macrourids and skates. In their deepest habitat (>1000 m depths), toothfish became a typical opportunistic predator, feeding mainly on relatively small and inactive fishes, squids and prawn‐like crustaceans Acanthephyra pelagica and Thymops birsteini. Decrease in hunting activity with depth could be related to a specific adaptation to keep neutral buoyancy by increase of lipid content in white muscles of D. eleginoides with size.