Tagging of Atlantic cod (Gadus morhua) with intragastric transmitters: effects of forced insertion and voluntary ingestion on retention, food consumption and survival

The effects of intragastric tagging on transmitter retention, food consumption and survival were investigated in Atlantic cod ( Gadus morhua L.) under laboratory conditions. Cod (60–84 cm) were intragastrically tagged with two different sizes of dummy acoustic transmitters (16 × 45 and 16 × 82 mm) using the methods of forced insertion and voluntary ingestion. Cod that were tagged by forced insertion began to regurgitate transmitters the first day after tagging and continued to do so throughout the period of study. The percentage of fish retaining transmitters in this group dropped to 50% after 32 days. By comparison, cod that were tagged by voluntary ingestion exhibited a 44-day delay in the onset of initial regurgitation. This was followed by a rapid rate of regurgitation, with the percentage of fish retaining transmitters in this group dropping to 50% after 60 days. These retention times are prolonged in comparison to previous studies and have been attributed to the lower temperature regime used in this study. Transmitter size did not significantly affect the duration of retention. During the first 20 days following tagging, the average food consumption per fish was higher in the voluntary ingestion group compared with the forced insertion and sham groups. Mortality levels were 13, 44 and 56% for the voluntary ingestion, forced insertion and sham groups, respectively, over the period of study. We associate low food consumption and high mortality rates in the first 20 days of the experiment with high levels of stress due to handling. The findings indicate that voluntary ingestion is a more viable technique for the intragastric tagging of cod in field studies.     

Behaviour of the abyssal grenadier, Coryphaenoides yaquinae, monitored using ingestible acoustic transmitters in the Pacific Ocean

Scavenging grenadier fishes, Coryphaenoides yaquinae , were tracked with ingestible acoustic tags at 5800 m depth in the central North Pacific. A free-fall vehicle comprising a video camera and recording system together with a hydrophone, ultrasonic acoustic receiver and recorder was deployed on the sea bed. Special acoustic transmitters (75 kHz) were wrapped in bait and attached below and in view of the camera. Grenadiers were observed to ingest the baited transmitters and move away to ranges of over 100 m within 5h, some returning several times to within range of the receiving system. Voluntary ingestion of transmitters by this grenadier demonstrates that an inacessible population can be studied and provides a basis for development of automated tracking systems for long-term monitoring of individual deep-sea fish.     

Sampling estuarine fish using multi-mesh gill nets: Effects of panel length and soak and setting times

Experiments on the effects of soak (1, 3 and 6 h) and setting times (18:00, 22:00 and 3:00 h), and panel length (20, 50, 120 m) on catches of fish in multi-mesh (38, 54 and 89 mm) gill nets were done in a south-east Australian estuary to develop an optimal, representative and standardized sampling methodology for future fishery-independent surveys of estuarine icthyofauna. Univariate analysis revealed that while the longest soak times and panels often caught significantly more numbers and species of fish, there were no differences between short and intermediate soak times. Further, where differences between soak time and panel length treatments were detected, fish were often being caught in the same relative proportions. Standardized catch-per-unit-of-effort (CPUE) (numbers of fish caught 20 m^− 1 of net h^− 1) decreased significantly with increasing soak time, but there were no differences in CPUE between different panel lengths for the total numbers of fish and key species caught. Multivariate analyses failed to detect any differences in fish assemblages between soak times and panel lengths for 38 and 89 mm mesh. However, inconsistent and species-specific differences in the abundances of common species accounted for differences in assemblage structure for the 54 mm mesh. Similarly, while the size ranges of most species of economic importance were comparable between different panel lengths and soak times, there were inconsistent differences in the proportions of fish captured across size classes for some species. Setting time had no effect on the mean numbers of fish or species caught, structure of assemblages or size-ranges of most species investigated. Based on these results, the use of 20-m panels soaked for 1 h at any time during the night was considered optimal for future surveys. The benefits of this uniform sampling methodology are discussed in terms of increased replication, reduced costs and the potential for lower fish mortality.     

Movements and Site Fidelity of two Juvenile Fish Species Using Surf Zone Nursery Habitats Along The Southeastern North Carolina Coast

We examined the extent of movements of juvenile Florida pompano, Trachinotus carolinus, and gulf kingfish, Menticirrhus littoralis, along an open ocean beach. Fishes were collected by seine at three sites along Masonboro Island and Carolina Beach, NC between 7 June and 7 July 1995. All specimens 40 mm standard length (SL) were tagged with coded wire tags and released at the capture sites. Between 7 July and 9 August and on 15–16 September Masonboro Island and northern Carolina Beach were surveyed for tagged fish. A controlled tag mortality/retention study was conducted for both species. Overall, 1569 Florida pompano (40–135 mm SL) were tagged. Sixty-one (3.9%) of these were recaptured, and only eight moved away from the original tagging sites. The largest movements by two Florida pompano were 2.1 and 10.5 km. Many fish remained at their original tagging sites for 21–27 days. Of 488 gulf kingfish (36–158 mm SL) tagged, 16 (3.3%) were recaptured. Gulf kingfish also exhibited little movement away from tagging sites during the study, with individuals remaining at original tagging sites up to 21 days. Stock size estimates for Florida pompano ranged from 3354 to 4670 among the tagging sites, with densities ranging from 1.9 to 2.6 fish m^–2. The remarkable site fidelity exhibited by these two species suggests that resources were not limiting or that predation pressure was not high enough to cause large scale movements during the study. This implies that local disturbances could impact behavior or survival of juvenile fishes in the surf zone.     

Prehistoric versus modern Baltic Sea cod fisheries: selectivity across the millennia

Combining Stone Age and modern data provides unique insights for management, extending beyond contemporary problems and shifting baselines. Using fish chronometric parts, we compared demographic characteristics of exploited cod populations from the Neolithic Period (4500 BP) to the modern highly exploited fishery in the central Baltic Sea. We found that Neolithic cod were larger (mean 56.4 cm, 95% confidence interval (CI)+/-0.9) than modern fish (weighted mean length in catch =49.5+/-0.2 cm in 1995, 48.2+/-0.2 cm in 2003), and older (mean ages=4.7+/-0.11, 3.1+/-0.02 and 3.6+/-0.02 years for Neolithic, 1995, and 2003 fisheries, respectively). Fishery-independent surveys in 1995 and 2003 show that mean sizes in the stock are 16-17 cm smaller than reflected in the fishery, and mean ages approximately 1-1.5 years younger. Modelled von Bertalanffy growth and back-calculated lengths indicated that Neolithic cod grew to smaller asymptotic lengths, but were larger at younger ages, implying rapid early growth. Very small Neolithic cod were absent and large individuals were rare as in modern times. This could be owing to selective harvests, the absence of small and large fish in the area or a combination. Comparing modern and prehistoric times, fishery selection is evident, but apparently not as great as in the North Atlantic proper.     

Diurnal patterns in the spatial relationships between saithe, Pollachius virens, schooling in the wild

Six saithe, length 35 to 43 cm, were tagged with acoustic transmitters and individuals tracked for between 9 and 508 h during May 1989. The tagged fish were members of a schooling group and during the study, pairs of tagged fish were tracked simultaneously for periods ranging from 9 to 139 h. These data have been used to investigate the spatial relationships between individuals schooling in the wild. Distances between individuals showed a distinct diel pattern. Tagged fish were generally closer together during the day, than either at night or the periods of dawn and dusk. Those circumstances under which pairs of tagged fish could be shown to school together were investigated by analysing the differences in their swimming directions. Schooling was generally evident during the hours of daylight, although the degree to which fish schooled varied between pairs. The degree to which the fish schooled was also dependent upon distance between the tagged individuals. Schooling could be demonstrated for saithe within 20 m of each other during the day and within 10 m in two out of three pairs at night.     

Validation of the water offloading technique for diet assessment: an experimental study with Cory’s shearwaters ( Calonectris diomedea )

We examined the effect of prey type, repeated stomach flushing, digestion time, and meal size on the assessment of dietary intake of captive adult Cory’s shearwaters (Calonectris diomedea). For each of Cory’s shearwaters’ main prey type (fish, cephalopod, and crustacea), we used three different meal sizes and four digestion times, stomach-flushing the birds 1, 4, 8, or 16 h after feeding. On average, fish and cephalopods showed similar percentages of mass recovery (between 23% and 33%), whereas crustaceans showed a recovery about 10–15% greater. Conversely, fish and crustaceans showed similar percentages of items recovered (between 52% and 77%), whereas cephalopods showed about 10–35% greater recovery rates. We found no significant differences in the percentage of individual prey items recovered and the interval between ingestion and recovery, over intervals ranging from 1 to 16 h.L.R. Monteiro was tragically killed in a plane crash in the Azores in December 1999.     

Patterns of movement and habitat use by leafy seadragons tracked ultrasonically

Nine adult leafy seadragons Phycodurus eques were tracked using ultrasonic telemetry for between 2-10 days around West Island, Australia. All fish except one moved within well-defined home ranges of up to 5 ha (using minimum convex polygon method). Short bursts of movement (at average velocities of 2-17 m h⁻¹) punctuated long periods (up to 68 h) without movement. The exceptional fish moved almost in a straight line away from its tagging location near the end of the tracking period, at a maximum velocity of 146 m h⁻¹. There was no constant diel pattern in movements; some fish moved more at night, others during the day. The time leafy seadragons spent over particular habitats compared to the area of those habitats available at the study site was greater for Posidonia seagrass, about as expected for kelp-covered reefs and bare sand patches, and less than expected for Amphibolis seagrass and boulders covered with brown algae. In searching for tagging effects, a comparison of movement immediately after tagging showed no difference with subsequent movements for most fish. The lack of tagging effect may be because the transmitter can be attached to the bony appendages away from the body of the fish. There was no sign of damage to fish upon removal of transmitters after tracking.     

Retention of intragastric transmitters after voluntary ingestion by captive cod, Gadus morhua L.

Squid-encapsulated acoustic transmitters were ingested by Atlantic cod, Gadus morhua , (3128–10 221 g) held in captivity at 12° C. Retention times ranged from 5 to > 21 days. There was no significant change in feeding rate after transmitter ingestion.     

Morphological changes during metamorphosis in cod (Gadus morhua L.), with particular reference to the development of the stomach and pyloric caeca

Metamorphosis in cod, Gadus morhua L., was investigated with respect to morphological changes in the alimentary tract (mouth, stomach and pyloric caeca) and external characters (finfold and fin development). Morphometric measurements and histological investigations were made on both larvae and juveniles and all fish lengths are given as standard length. Median unpaired fins started to develop when the fish were 9–10 mm in length. The larval median finfold was resorbed when fish reached a size of 11–15 mm. The vertebrae were formed in fish of 11–20 mm. The supraterminal mouth, seen in larvae <12 mm, changed gradually to a sub-terminal position in juveniles larger than 65 mm. The development of the stomach and pyloric caeca started at 15 mm and the juvenile form was seen in fish of 40 mm. The ability to store and grind food particles in the stomach developed gradually as fish increased in size from 20 to 40 mm. Compared to many other teleost species, cod juveniles are relatively large when the stomach and pyloric caeca become fully developed. These findings may have important implications for both start-feeding of cod larvae and the weaning of juveniles on to artificial diets.     

Scaling effects on the neuromuscular system,twitch kinetics and morphometrics of the cod,Gadus morhua

Size related changes in muscle twitch kinetics, morphometrics and innervation have been examined in cod, Gadus morhua. Fish size ranged from 9 cm to 45 cm in total length (L).

Twitch contraction time (time to 90% relaxation), scaled in proportion to L^0.29. Scaling of morphometric parameters was essentially geometric. Mean cross‐sectional area and weight of the myotomal muscle scaled in proportion to L^2.05 and L^3.08 respectively. These results are discussed in the light of alterations in length specific swimming performance and kinematics as fish grow.

During growth, the number of endplates terminating on each fast fibre increased, from around 10 on fibres 2 mm in length (～10 cm fish) to 20 on 10 mm fibres (—40 cm fish). However, mean spacing between endplates increased from around 0.25 mm to 0.50 mm. The functional significance of polyneuronal innervation in teleost fast muscle fibres is discussed.     

Comparison between PIT and radio telemetry to evaluate winter habitat use and activity patterns of juvenile Atlantic salmon and brown trout

Winter habitat use and activity patterns of juvenile Atlantic salmon and brown trout were analysed in a comparative study between Passive Integrated Transponder (PIT) technology, radio telemetry and underwater observation by snorkelling. Two study periods were conducted in Stoney River, Newfoundland, Canada. During Study period I, 49 juvenile Atlantic salmon (fork length: 11.0–18.0 cm) and 7 brown trout (11.0–17.3 cm) were tagged with PIT tags and/or radio transmitters in late winter of 2004. During Study period II, 18 juvenile Atlantic salmon (fork length: 12.0–18.4 cm) and 23 brown trout (10.9–20.8 cm) were tagged and tracked twice a day at 10:00 h and 22:00 h on five consecutive days in late winter of 2005. From the 56 fish released during Study period I, on average 19.6 ± 6.0% of the PIT tagged fish and 99.3 ± 2.2% of the radio tagged fish were relocated during any given survey. Over the Study period II, 39% of fish emigrated from the study site. PIT technology had an efficiency of 39.2 ± 14.1% to detect the remaining fish. In contrast, radio telemetry relocated on average 96.9 ± 6.5% of the tagged fish whereas by snorkelling on average only 4.1 ± 5.6% of the tagged fish were observed. PIT telemetry may however be more efficient in smaller, less heterogeneous streams. The advantage of PIT technology over radio telemetry is clearly that it is relatively less costly permitting higher numbers of individuals to be tagged and there is no limit in the operational life of the transponder. In winter, juvenile salmonids preferred low flow velocity and no preferences were observed for any specific water depth over the range of available water depths. Fish selected preferentially boulder habitat over other substrates in the environment. Habitat utilisation did not differ between day and night. The use of winter preference indices may be important for future habitat modelling.     

A field and experimental evaluation of the effect of data storage tags on the growth of cod

The effect of electronic data storage tags (DSTs) on the growth of cod Gadus morhua was evaluated in laboratory and field experiments. In the first laboratory experiment, large DSTs (60 × 18 mm, 3 g in water ) attached externally for 3 months did not have any effect ( anova , P  > 0·05) on the growth of adult cod (mean ±  s . d . 65 ± 4·5 cm total length) relative to untagged adult cod. In a second experiment, small DSTs (34 × 11 mm, 1·5 g in water) implanted into young cod (48·1 ± 4·4 cm) for an 8 month period did not have any effect upon the growth relative to untagged controls ( anova , P  > 0·05). Length data returned from tagging experiments conducted on adult cod (57·3 ± 7·5 cm) in the North Sea showed that the growth of fish tagged either externally or internally with large DSTs was not different ( t ‐test, P  > 0·05). Attachment wounds, however, provided evidence that external attachment of DSTs should be avoided unless sensor configuration requires access to the external environment, and that internal implantation should be preferred whenever possible.     

Ontogeny of haemoglobin polymorphism in cod

Haemoglobins of 0-group cod from 30 to 150 mm length were analysed by agar gel electrophoresis and isoelectric focusing to identify the lower size limit for determination of the haemoglobin polymorphisms in cod; both the main system which has been known for more than 30 years and the recently described subgroups. The juvenile fish studied had higher concentration of the HbII-component (split into three bands at these fish sizes) compared to older fish. The main haemoglobin types could be distinguished clearly and reliably at a fish size of about 35 mm and above. The subgroups of the HbI-components, however, could not be identified reliably until the cod were about 80 mm.     

Growth and mortality of exploited fishes in China's coastal seas and their uses for yield‐per‐recruit analyses

Given the scarcity of information suitable for other forms of fish stock assessments, the growth, and mortality of 10 important marine exploited fishes in China's coastal seas were estimated, based on published length‐frequency data and the ELEFAN approach and software. The resulting parameters, complemented with growth curves from FishBase were then used for yield‐per‐recruit analyses. These 10 species were both traditional commercial species (Trichiurus lepturus, Larimichthys polyactis, Scomber japonicus) and less commercial interest species (Coilia nasus, Engraulis japonicus, Decapterus maruadsi, Psenopsis anomala, Coilia mystus, Setipinna tenuifilis, Harpadon nehereus). The yield‐per‐recruit analyses indicated that the fishing mortality experienced by these 10 fish species was very high, especially in view of the extremely small cod‐end mesh sizes deployed by Chinese trawlers, of about 10 mm or less (vs. 25 mm as minimum legal size). More precisely, all 10 species suffered from growth overfishing, which was very pronounced in large species. Indeed, enlarging cod‐end mesh size, by increasing mean sizes at first capture, would increase yield‐per‐recruit for all 10 fish species by a factor of up to 2.5. Obviously, such increases in mesh size would have to be in tandem with other management measures designed to ensure the sustainability of China's marine fisheries resources.     

The combined use of acoustic tracking and echosounding to investigate the movement and distribution of common bream (Abramis brama) in the River Trent,England

Spatial behaviour and distribution of fishes along a 7.6-km lowland reach of the River Trent, England, were examined using two complementary telemetry techniques: acoustic tracking to assess the movement and activity of common bream Abramis brama (L.) and quantitative echosounding for measuring the density and distribution of fish shoals. Nine adult bream (39.3–53.2 cm) were tracked by means of intraperitoneally implanted acoustic transmitters from 19 July to 12 September 2000. Home range size varied between 0.35 and 5.40 km of river length over this period. Bream were relatively inactive during daylight hours, began moving near dusk, and tended to move throughout the night. A distinct daytime residence area was occupied by most tagged fish on most occasions, while river use at night was more variable between individuals. Mobile echosounding surveys, with the transducer beaming horizontally across the river, conducted at night between July and September 2000, showed a highly contagious fish distribution within the study reach. For 200-m sections of river, there was a negative correlation between the relative frequency of acoustic tracking fixes at night and mean fish densities, as measured by echosounding for targets larger than –50 dB (c. 5-cm long). However, there was a highly significant positive rank correlation between the relative frequency of acoustic tracking fixes and acoustic targets larger than –30 dB (c. 22-cm long), most of which in this river are bream. This suggests that telemetry and echosounding can, in this part of the River Trent, be combined to provide valuable spatial information at individual and population scales for bream.     

Effect of position-fixing interval on estimated swimming speed and movement pattern of fish tracked with a stationary positioning system

Ultrasonic telemetry using stationary positioning systems allows several fish to be tracked simultaneously, but systems that are incapable of sampling multiple frequencies simultaneously can record data from only one transmitter (individual) at a time. Tracking several individuals simultaneously thus results in longer intervals between successive position fixes for each fish. This deficiency leads to loss of detail in the tracking data collected, and may be expected to cause loss of accuracy in estimates of the swimming speeds and movement patterns of the fish tracked. Even systems that track fish on multiple frequencies are not capable of continuous tracking due to technical issues. We determined the swimming speed, area occupied, activity rhythm and movement pattern of cod (Gadus morhua) using a stationary single-channel positioning system, and analysed how estimates of these behavioural parameters were affected by the interval between successive position fixes. Single fish were tracked at a time, and position fixes were eliminated at regular intervals in the original data to generate new data sets, as if they had been collected in the course of tracking several fish (2–16). In comparison with the complete set, these data sets gave 30–70% decreases in estimates of swimming speed depending on the number of fish supposedly being tracked. These results were similar for two individuals of different size and activity level, indicating that they can be employed as correction factors to partly compensate for underestimates of swimming speed when several fish are tracked simultaneously. Tracking `several' fish only slightly affected the estimates of area occupied (1–15%). The diurnal activity rhythm was also similar between the data sets, whereas details in search pattern were not seen when several fish were tracked simultaneously.     

Movements of Atlantic salmon, Salmo salar L., in an estuary in south-west England

In the years 1985–1987, salmon were tracked in the estuary of the River Fowey, Cornwall using the 'sonar buoy' system developed at the M.A.F.F. Fisheries laboratory and described by Solomon & Potter (1988). Sixty-one salmon were caught and tagged in the estuary, 25 with radio and 36 with combined radio and acoustic transmitters. About half these fish were not recorded entering the freshwater reaches of the R. Fowey, and from tag recoveries it seems likely that about 27% were returning to other rivers. For fish returning to the R. Fowey the time between tagging and entry into fresh water varied between 9 h and 130 days. In the estuary the fish largely moved in the same direction as the tidal currents, although movements over the ground against the current were more common in the bottom 2 km and the top 2 km. During periods of low river-flow in summer all salmon tracked in the estuary dropped back out to sea where they stayed for varying periods: some of these made several sorties past the estuary mouth and on occasions well up the estuary before finally entering fresh water. During periods of higher river discharge, in both the summer and the autumn, some fish remained in the estuary for longer periods, sometimes holding position in deep, sheltered water. Most fish entered fresh water at night during periods of increased freshwater discharge, although at very high flows fish also entered in daylight.     

Hypoxia tolerance in Atlantic cod

Oxygen saturation levels that killed 50 and 5% of cod Gadus morhua over 96 h averaged 21·2 and 27·7%, respectively. No fish survived at 10% saturation and only a few survived at 16% saturation, whereas no mortality occurred at 34 and 40% oxygen saturation. Since metabolic rate and oxygen consumption increase with increasing temperature, we hypothesized that cod would be less tolerant to hypoxic conditions at 6 than at 2° C. However, temperature (2 and 6° C) had no measurable impact on cod survival. Small (mean & S.D.; 45·2 ± 4·2 cm) and large (57·5 ± 3·8 cm) cod had the same tolerance to hypoxia. At the end of the experiments, hypoxia had a significant effect on blood haematocrit, mean cellular haemoglobin content, liver lactate, plasma glucose and plasma lactate, but accounted for only a small fraction (< 10%) of the variation, except for plasma lactate which exhibited a strong response with concentrations increasing progressively with decreasing levels of oxygen saturation. Temperature had a significant effect on most variates in normoxia and hypoxia. Variates also affected by oxygen level showed significant interactions between oxygen and size or temperature effects. However, these interactions accounted for only a small proportion of the variation. Physiological parameters indicated that extending the duration of our tests beyond 96 h would not have changed our estimates of the lethal thresholds. Hypoxic conditions are a permanent feature of the deep waters of the Gulf of St Lawrence. This study shows that a significant portion of the benthic habitats in the Gulf are uninhabitable for cod which would be expected to avoid waters below 28% oxygen saturation.     

Factors influencing the behavior and duration of residence of adult Chinook salmon in a stratified estuary

The duration of residence, behavior, and thermal experience of adult Chinook salmon (Oncorhynchus tshawytscha) in a stratified lagoon-type estuary over a four-year period was determined using acoustic and radio biotelemetry transmitters and archival temperature tags. Results did not support the hypothesis that adult Chinook salmon would hold extensively and migrate slowly through the estuarine lagoon with return trips to the sea and use of the salt wedge for behavioral thermo-osmoregulation in response to high summer water temperatures. Passive tidal transport was not observed as predicted and entry from the sea to the estuarine lagoon occurred during all tidal phases suggesting that conserving energy was not a priority for fish during the estuarine phase of their migration. An unexpected finding was that the persistent pursuit pressure of pinnipeds, especially from California sea lions (Zalophus californianus), appeared to negate the suitability of estuarine lagoon as holding habitat. This finding led to the hypothesis that the risk of pinniped predation was primarily responsible for the brief residence (<24 h on average) and rapid migration (mean 1.2 km/h, 0.42 body lengths/s) of adult Chinook salmon in the estuarine lagoon, especially given its relatively small size (7 km). This hypothesis will be difficult to test without exclusion of pinnipeds. Thermal records of fish that retreated back to the sea after tagging suggested that use of the marine river plume could be important for osmotic adaption to allow rapid migration through estuaries, which often contain concentrations of predators anticipating the return of migrating salmon. Alternatively, use of the marine river plume could have occurred independent of predators. Fish retreating back to the sea should be considered a tagging effect as downstream movement commonly occurs when adult salmonids are released after tagging. There was no evidence of any other tagging effects or biased behavior by fish tagged in this study. Regardless of the reasons, the successful migration of fish that held for weeks in the sea after tagging indicates that there could be considerable flexibility in the river entry timing of some salmon stocks. Tagging adult salmonids in the sea as they approach their natal rivers is ideal but the associated logistical challenges and expense make tagging fish immediately upon entrance to river mouths the next best option when possible.