A hydro-mechanical approach to the scaling of swimming performance in the sand flathead Platycephalus bassensis Cuvier: effects of changes in morphological features based on fish size

The swimming performance of Platycephalus bassensis at steady speed was assessed with an emphasis on hydrodynamics. The minimum swimming speed to maintain hydrostatic equilibrium for P. bassensis of 0·271 m total length ( L _T) was calculated to be 1·06 L _T s⁻¹. At this speed, the required lift to support the mass of the fish was equivalent to 6·6% of the fish mass; 82·7% of which was created by the body as a hydrofoil, and the rest of which was created by the pelvic fins as hydrofoils. The minimum swimming speed decreased with the L _T of the fish and ranged from 1·15 L _T s⁻¹ for a fish of 0·209 m to 0·89 L _T s⁻¹ for a fish of 0·407 m. The forward movement per tail-beat cycle ( i.e. stride length) was described with an equation including quantities of morphological and hydro-mechanical relevance. This equation explained that stride length was increased by the effect of turbulence characterized by the Reynolds number and demonstrated the morphological and hydro-mechanical functional design of the fish for maximizing thrust and minimizing drag. The larger span of the caudal fin and caudal tail-beat amplitude was associated with larger stride length, whereas greater frictional drag was associated with smaller stride length.     

Age and growth of Anguilla anguilla in the Camargue lagoons

Age and total length ( L _T) data from a 11 year monitoring of the Anguilla anguilla eel population of the Camargue lagoons (Rhône delta, southern France) were collected for glass, yellow and silver eels. Three distinct models were calibrated to describe the growth process of undifferentiated eels, females and males, respectively. Uncertainty of parameter estimates was evaluated by bootstrapping. Females were characterized by larger asymptotic body size ( L _T) than males (580 ± 50 v . 388 ± 13 mm) and faster growth, whilst the Brody growth coefficient was larger for males than for females (means ±  s . d . 3·00 10⁻³ ± 1·68 10⁻³ v . 1·73 10⁻³ ± 0·50 10⁻³). Sexual differentiation was estimated to begin at 204 ± 38 mm mean ±  s . d ., i.e . at the end of the second year in the lagoons, well before the L _T at which macroscopic differentiation became possible ( c . 300 mm). Males probably leave the lagoon or die (due to either natural or fishing mortality) within the first 3 years, whilst females can remain up to 5 years. Sexual differentiation and maturation have a major role in shaping the L _T structure of the population. The L _T and mass ( M ) data were fitted by allometric curves     . The calibration of distinct curves for data from different years indicated that the allometric coefficient a was subject to wider interannual fluctuations than the allometric exponent b . A negative correlation linked the average L _T and the allometric exponent ( r  = −0·58, P  < 0·01).     

Early growth and juvenile population structure of lemon sharks Negaprion brevirostris in the Atol das Rocas Biological Reserve, off north‐east Brazil

Lemon sharks Negaprion brevirostris were sampled in the Atol das Rocas, a nursery area, on nine occasions from March 1999 to October 2003, during which 157 individuals were tagged and 35 were recaptured. The male : female sex ratio of captured individuals was 1 : 1·12. Mean ±  s . d . growth rates were 24·7 ± 3·4 cm year⁻¹ in total length ( L _T), 20·7 ± 3·2 cm year⁻¹ in fork length, and 19·5 ± 2·7 cm year⁻¹ in precaudal length. There was no significant difference in growth rates between males and females. Mean ±  s . d . increase in mass was 2565 ± 762 g year⁻¹. The von Bertalanffy growth parameters estimated by the Fabens method based on L _T were: k  = 0·077, L _∞ = 399·9 cm and t ₀ = −2·16. Despite the large variation of environmental conditions, particularly of tidal range and currents, and the lack of protective mangrove cover in the nursery area at Atol das Rocas, juvenile lemon sharks grew relatively faster than at other nurseries. Such rapid growth could be a response to abundant food availability or high risk of predation by adults that enter the nursery area.     

A histological study on the development of the digestive system of Pseudosciaena crocea larvae and juveniles

The ontogenetic development of the gut and accessory organs in large yellow croaker Pseudosciaena crocea was investigated using light microscopy from hatching up to the juvenile stage (40 days post hatch, dph). At 3 dph (mean ±  s . d ., 4·1 ± 0·1 mm total length, L _T), coinciding with the buccopharynx opening, larvae started to feed exogenously, and the gut consisted of a well‐developed buccopharynx, a partially‐differentiated oesophagus and an intestine divided in three regions (anterior intestine, intermediate intestine and rectum). Yolk reserves were not completely depleted at the onset of exogenous feeding, and a period of mixed nutrition was observed up to 6 dph (4·3 ± 0·1 mm L _T), when yolk was definitively exhausted. Important morphological changes occurred at the end of the larval period, coinciding with metamorphosis. At 17 dph (6·8 ± 0·6 mm L _T), pyloric caeca differentiated at the junction of the pyloric stomach and the anterior intestine. Gastric glands were first observed at 21 dph (9·2 ± 1·2 mm L _T), coinciding with the morphological development of the stomach in three different regions (cardiac, fundic and pyloric) according to the histological characteristics of their mucosa. At this age, large longitudinal folds appeared in the median and posterior oesophageal mucosa. These morphological and histological features suggested the achievement of a digestive system characteristic of large yellow croaker juveniles and adults.     

Mackerel icefish size and age differences and long‐term change at South Georgia and Shag Rocks

This study analysed the total length ( L _T)‐frequency distribution of mackerel icefish Champsocephalus gunnari at South Georgia and Shag Rocks from nine bottom trawl surveys at South Georgia and eight at Shag Rocks between 1987 and 2002. The estimated mean L _T of age‐classes 1+, 2+, 3+ and 4+ years during January were, respectively, 14·7, 23·5, 29·8 and 35·1 cm at South Georgia. Age‐classes 1+, 2+ and 3+ years were 18·3, 26·2 and 33·8 cm at Shag Rocks. The derived Bertalanffy growth parameters for South Georgia were: L _∞ = 51·7 cm, k  = 0·27 and t ₀ = −0·26. The mean L _T of each age‐class of C. gunnari at Shag Rocks was significantly larger than at South Georgia, equivalent to c . 5 months growth, although the annual growth in L _T was similar. This is further evidence that C. gunnari hatched earlier at Shag Rocks. At South Georgia, the mean L _T of age‐classes 1+ and 3+ years were correlated, and significantly decreased between 1987 and 2002, and were smaller following warmer summers. This decrease in the size of C. gunnari may be the result of reduced food availability linked to climate warming.     

Reproductive biology of the milk shark Rhizoprionodon acutus and the bigeye houndshark Iago omanensis in the coastal waters of Oman

Female Rhizoprionodon acutus were found to mature between 62 and 74 cm total length ( L _T) whereas males matured between 63 and 71 cm L _T. The L _T at 50% maturity was 64·3 cm for females and 64·7 cm for males. Litter size varied from one to six embryos, and was positively correlated with maternal L _T. Female embryos outnumbered males by a ratio of 2·3:1. The size at birth was c . 37 cm L _T. Full‐term embryos and post‐partum females were observed during all seasons although their occurrence was highest in spring. Spermatozoa were rarely recorded in the oviducal gland, indicating that this species does not store sperm. It was not possible to generate maturity curves for Iago omanensis but it was evident that females matured by the time they reached 35 cm and males were mature by 31 cm L _T. This species displayed a clearly defined reproductive cycle with parturition occurring primarily in spring, after a gestation period of c . 1 year. Maximal embryo size was 19 cm L _T while maximal litter size was 24 embryos. The oviducal gland appeared to act as a seminal receptacle and it appeared that females may utilize these stores by not mating every year.     

Endurance swimming of diploid and triploid Atlantic salmon

When groups of diploid (mean ±  s . e . fork length, L _F) 33·0 ± 1·4 cm and triploid (35·3 ± 0·5 cm) Atlantic salmon Salmo salar were forced to swim at controlled speeds in a carefully monitored 10 m diameter 'annular' tank no significant difference was found between the maximum sustained swimming speeds ( U _ms, maintainable for 200 min) where the fish swam at the limit of their aerobic capability. Diploids achieved 2·99 body lengths per second (bl s⁻¹)(0·96 m s⁻¹) and triploids sustained 2·91 bl s⁻¹(1·02 m s⁻¹). The selection of fish for the trials was based on their ability to swim with a moving pattern projected from a gantry rotating at the radius of the tank and the selection procedure did not prove to be significant by ploidy. A significant difference was found between the anaerobic capabilities of the fish measured as endurance times at their prolonged swimming speeds. During the course of the experimentation the voluntary swimming speed selected by the fish increased and the schooling behaviour improved. The effect of the curvature of the tank on the fish speeds was calculated (removing the curved effect of the tank increased the speed in either ploidy by 5·5%). Implications of the endurance times and speeds are discussed with reference to the aquaculture of triploid Atlantic salmon.     

Some morphometric and biochemical features of ready-to-migrate silver and pre-migratory yellow stages of the shortfin eel of south-eastern Australian waters

The mean total length ( L _T), mass and age of ready to migrate female silver shortfin eels Anguilla australis from the Hopkins River estuary and the mouth of the Merri River in south-eastern Australia, were 83·2 ± 1·2 cm, 1051 ± 51 g, and 17·2 ± 1·79 years, respectively. The eye index ( I _E) of the silver shortfin eels was < 5·2 (mean 7·64 ± 0·29) and differed significantly from that of the yellow shortfin eels collected from two other sites. The I _E increased with L _T (mm) and was related by log I _E= 2·656 log L _T6·925. The per cent moisture, protein and ash content of the liver of silver shortfin eels was significantly lower than in yellow shortfin eels, but lipid content was significantly higher in the former (35·5 ± 2·0%). The mean mass μg mg lipid ⁻) of saturates (230·4 ± 2·6 v . 181·7 ±2·6), monoenes (367·4 ± 6·3 v . 290·8 ± 8·9) and PUFA (177·3 ± 5·3 v . 159·7 ± 4·6) in muscle was significantly higher, and the great majority of individual fatty acids was found also in higher quantities in silver shortfin eels. In the liver, the PUFA found in the highest quantity was 22:6n-3, except in shortfin eels from Hopkins River estuary, and the amount of 18:2n-6 in the liver of silver shortfin eels was significantly higher than that in yellow shortfin eels but the reverse was true of 20:4n-6. In both muscle and liver tissues the saturate 16:0 and the monoene 18:ln-9 collectively accounted for >50% of all the fatty acids in the lipid.     

Life‐history variation among local populations of Atlantic cod from the Norwegian Skagerrak coast

Small‐scale spatial variation in life history was found among genetically distinct local populations of Atlantic cod Gadus morhua from the Norwegian Skagerrak coast. Among populations, age at 50% maturity varied from 2·6 to 3·8 years, total ( L _T) length at 50% maturity from 35 to 60 cm, annual survival from 33 to 64%, mean L _T at age 4 years from 43 to 63 cm, and mean backcalculated L _T at age 1 year from 8 to 12 cm.     

Reproductive strategy of a primitive temperate notothenioid Eleginops maclovinus

The reproductive biology of Eleginops maclovinus was examined in the Falkland Islands between October 2000 and December 2002. Males predominated at total lengths ( L _T) of 10 to 52 cm and females at >53 cm L _T. Length frequency analysis showed a bimodal distribution with females representing the larger mode for every month during the study period. Gonad histology revealed that 19% of the histological samples studied were considered to be those from hermaphrodites: morphologically as male gonads but containing protoplasmic oocytes. It was therefore concluded that E. maclovinus is a protandrous hermaphrodite. The size of first (male) maturation of E. maclovinus was 30·73 cm L _T. Males and females matured from August onwards and spawning occurred between September and December at depths of >40 m. Eleginops maclovinus has the smallest eggs and highest fecundity among the notothenioids. The highest potential fecundity was attained at maturity stage III with c . 48 million eggs. Because of further oocyte resorption, this value gradually decreased until the final fecundity ranged from 1·1 to 7·3 million eggs. Oocyte length frequencies in ovaries suggested that E. maclovinus was a batch spawner. Hydrated oocytes were found to contain a large perivitelline space indicating that the egg had a pelagic development.     

Piscivory and trophic position of Anguilla anguilla in two lakes: importance of macrozoobenthos density

The feeding habits of the European eel Anguilla anguilla (>300 mm total length, L _T) were compared in two lakes of different environmental state: Lake Großer Vätersee (LGV), Germany (clear water, mesotrophic and submerged macrophytes), and Lake Vallum (LV), Denmark (turbid, eutrophic and no submerged macrophytes). The density of macrozoobenthos was higher in LV (3500 individuals m⁻²) than in LGV (1500 individuals m⁻²). The abundance of small prey fishes (40–99 mm L _T) was highest in LV. In LV, A. anguilla fed on macrozoobenthos, in particular, chironomid larvae. In LGV, A. anguilla used fishes as the main food component. Stable isotope analyses confirmed the stomach contents dietary results. The estimated mean ± s.d . trophic positions of A. anguilla in LGV (3·7 ± 0·2) was one level higher than those of fish in LV (2·7 ± 0·2). Based on these results, it is concluded that piscivory among A. anguilla was generally controlled by the density of macrozoobenthos. Stable isotope analysis further indicated that A. anguilla may act as integrators between benthic and pelagic food webs when density of insect larvae is low.     

The effect of external dummy transmitters on oxygen consumption and performance of swimming Atlantic cod

Decreased critical swimming speed and increased oxygen consumption (     ) was found for externally tagged Atlantic cod Gadus morhua swimming at a high speed of 0·9 body length (total length, L _T) s⁻¹. No difference was found in the standard metabolic rate, indicating that the higher     for tagged cod was due to drag force rather than increased costs to keep buoyancy.     

The angle of attack of the body of common bream while swimming at different speeds in a flume tank

The body attack angle of common bream Abramis brama varied with swimming speed and was best described by θ = −3·32 (±0·24) − 9·23 (±0·54)e^{− u} ( r ² = 0·56, P  < 0·0001; ±1 s . e . given in parentheses), where θ is the body attack angle and u is swimming speed. The hypothesis that neutrally buoyant fishes may swim with body attack angles deviating increasingly from 0° as the swimming speed decreases is supported.     

Is tilting behaviour at low swimming speeds unique to negatively buoyant fish? Observations on steelhead trout,Oncorhynchus mykiss,and bluegill,Lepomis macrochirus

When swimming at low speeds, steelhead trout and bluegill sunfish tilted the body at an angle to the mean swimming direction. Trout swam using continuous body/caudal fin undulation, with a positive (head-up) tilt angle ( 0 , degrees) that decreased with swimming speed ( u , cm s⁻¹) according to: 0 =(164±96).u^{(−1.14±0.41)} (regression coefficients; mean±2 s.e. ). Bluegill swimming gaits were more diverse and negative (head down) tilt angles were usual. Tilt angle was −3·0 ± 0.9° in pectoral fin swimming at speeds of approximately 0.2–1.7 body length s⁻¹ (Ls⁻¹; 3–24 cm s⁻¹), −4.5 ±2.6° during pectoral fin plus body/caudal fin swimming at 1·2–1·7 L s⁻¹ (17–24cm s⁻¹), and −5.0± 1.0° during continuous body/caudal fin swimming at 1.6 and 2.5 L s⁻¹ (22 and 35cm s⁻¹). At higher speeds, bluegill used burst-and-coast swimming for which the tilt angle was 0.1±0.6°. These observations suggest that tilting is a general phenomenon of low speed swimming at which stabilizers lose their effectiveness. Tilting is interpreted as an active compensatory mechanism associated with increased drag and concomitant increased propulsor velocities to provide better stabilizing forces. Increased drag associated with trimming also explains the well-known observation that the relationship between tail-beat frequency and swimming speed does not pass through the origin. Energy dissipated because of the drag increases at low swimming speeds is presumably smaller than that which would occur with unstable swimming.     

The reproduction, age and growth of the spotted rockling

Biology of the Macaronesian endemic rockling Gaidropsarus guttatus was studied in the Azores. The overall sex ratio from the samples was highly in favour of females (1 : 6·33). The growth parameters were L _∞ = 24·23, k  = 1·219 and t ₀ = −0·059. Fish matured at 15 cm L _T and the spawning season was strongly clustered in April.     

Physiological impact of sea lice on swimming performance of Atlantic salmon

Atlantic salmon Salmo salar were infected with two levels of sea lice Lepeophtheirus salmonis (0·13 ± 0·02 and 0·02 ± 0·00 sea lice g⁻¹). Once sea lice became adults, the ventral aorta of each fish was fitted with a Doppler cuff to measure cardiac output ( ̇ ), heart rate ( f _H) and stroke volume ( V _S) during swimming. Critical swimming speeds ( U _crit) of fish with higher sea lice numbers [2·1 ± 0·1 BL (body lengths) s⁻¹] were significantly lower ( P  < 0·05) than fish with lower numbers (2·4 ± 0·1 BL s⁻¹) and controls (sham infected, 2·6 ± 0·1 BL s⁻¹). After swimming, chloride levels in fish with higher sea lice numbers (184·4 ± 11·3 mmol l⁻¹) increased significantly (54%) from levels at rest and were significantly higher than fish with fewer lice (142·0 ± 3·7 mmol l⁻¹) or control fish (159·5 ± 3·5 mmol l⁻¹). The f _H of fish with more lice was 9% slower than the other two groups at U _crit. This decrease resulted in ̇ not increasing from resting levels. Sublethal infection by sea lice compromised the overall fitness of Atlantic salmon. The level of sea lice infection used in the present study was lower than has previously been reported to be detrimental to wild Atlantic salmon.     

Effect of delayed first feeding on development and feeding ability of Paralabrax maculatofasciatus larvae

The impact of delayed feeding early in development on late feeding ability and development of spotted sand bass Paralabrax maculatofasciatus larvae was examined. Larvae were sampled from hatching until day 19 after delayed feeding for zero (control), 1, 2, or 3 days. Feeding incidence was evaluated as the percentage of larvae with food in the gut and feeding intensity was measured by direct counting of prey after dissection of the gut. Delayed feeding effects due to starvation were observed early in development. By day 5, significant differences ( P  < 0·05) were observed between controls and larvae submitted to degrees of delayed feeding, including total length ( L _T), eye diameter, and the size of the head, liver, gut, muscles and body. Differences were still apparent in L _T, and body, liver and muscle heights at the end of the studied period. Larvae under total starvation did not survive beyond day 5. Initial feeding incidence was 35, 60, 90 and 10% for larvae first fed on day 2, 3, 4 or 5, respectively. Mean ±  s . d . feeding intensity was 3·6 ± 0·8, 2·8 ± 1·3, 5·2 ± 0·3, and 10·2 ± 1·5 rotifers per larva depending on whether larvae were initially fed on day 2, 3, 4 or 5, respectively. The point of no return occurred between 4 and 5 days after hatching.     

Energy savings in sea bass swimming in a school: measurements of tail beat frequency and oxygen consumption at different swimming speeds

Tail beat frequency of sea bass, Dicentrarchus labrax (L.) (23.5 ± 0·5 cm, L_T ), swimming at the front of a school was significantly higher than when swimming at the rear, for all water velocities tested from 14·8 to 32 cm s⁻¹. The logarithm of oxygen consumption rate, and the tail beat frequency of solitary swimming sea bass (28·8 ± 0·4 cm, L_T ), were each correlated linearly with swimming speed, and also with one another. The tail beat frequency of individual fish was 9–14% lower when at the rear of a school than when at the front, corresponding to a 9–23% reduction in oxygen consumption rate.     

Endurance swimming of European eel

A long‐term swim trial was performed with five female silver eels Anguilla anguilla of 0·8–1·0 kg ( c . 80 cm total length, L _T) swimming at 0·5 body lengths (BL) s⁻¹, corresponding to the mean swimming speed during spawning migration. The design of the Blazka‐type swim tunnel was significantly improved, and for the first time the flow pattern of a swim tunnel for fish was evaluated with the Laser‐Doppler method. The velocity profile over three different cross‐sections was determined. It was observed that 80% of the water velocity drop‐off occurred over a boundary layer of 20 mm. Therefore, swim velocity errors were negligible as the eels always swam outside this layer. The fish were able to swim continuously day and night during a period of 3 months in the swim tunnel through which fresh water at 19° C was passed. The oxygen consumption rates remained stable at 36·9 ± 2·9 mg O₂ kg⁻¹ h⁻¹ over the 3 months swimming period for all tested eels. The mean cost of transportation was 28·2 mg O₂ kg⁻¹ km⁻¹. From the total energy consumption the calculated decline in fat content was 30%. When extrapolating to 6000 km this would have been 60%, leaving only 40% of the total energy reserves for reproduction after arriving at the spawning site. Therefore low cost of transport combined with high fat content are crucial for the capacity of the eel to cross the Atlantic Ocean and reproduce.     

Variation in the diet of Genypterus blacodes(Ophidiidae) around the Falkland Islands

The diet change with size, season and area was investigated using the stomachs of 496 kingclip Genypterus blacodes collected around the Falkland Islands (south‐west Atlantic) between August 2001 and September 2002. The key prey species were rockcod Patagonotothen spp., benthic isopods and Patagonian grenadier Macruronus magellanicus . Kingclip <50 cm total length ( L _T) fed mainly on crustaceans and small fishes. With size the diet shifted away from crustaceans towards Patagonotothen spp. in kingclip 50–100 cm L _T, and finally towards larger fishes such as M. magellanicus and Micromesistius australis australis in kingclip >100 cm L _T. The niche breadth was highest in fish >100 cm L _T and the lowest in fish <50 cm L _T. The larger kingclip generally selected larger individuals of the same prey species, with the exception of the Patagonian squid Loligo gahi , where all ingested squid were of similar size, regardless of the predator length. The importance of the main prey species varied substantially between five consequent seasons studied, and appeared to follow the seasonal abundance and availability of prey. The spatial variability in the diet was found in kingclip caught in regions occupied by transformed temperate and sub‐Antarctic waters. The rockcod, which is available throughout the year around the Falkland Islands, was the most important prey in the kingclip diet. Kingclip takes advantage of other seasonally abundant prey species during their seasonal migrations ( e.g . L. gahi ) and also scavenge on discards from fishing vessels when available.