Behaviour that influences dispersal and connectivity in the small,young larvae of a reef fish

Determining the scale of larval dispersal and population connectivity in demersal fishes is a major challenge in marine ecology. Historically, considerations of larval dispersal have ignored the possible contributions of larval behaviour, but we show here that even young, small larvae have swimming, orientation and vertical positioning capabilities that can strongly influence dispersal outcomes. Using young (11–15 days), relatively poorly developed (8–10 mm), larvae of the pomacentrid damselfish, Amblyglyphidodon curacao (identified using mitochondrial DNA), we studied behaviour relevant to dispersal in the laboratory and sea on windward and leeward sides of Lizard Island, Great Barrier Reef. Behaviour varied little with size over the narrow size range examined. Critical speed was 27.5 ± 1.0 cm s⁻¹ (30.9 BL s⁻¹), and in situ speed was 13.6 ± 0.6 cm s⁻¹. Fastest individuals were 44.6 and 25.0 cm s⁻¹, for critical and in situ speeds, respectively. In situ speed was about 50% of critical speed and equalled mean current speed. Unfed larvae swam 172 ± 29 h at 8–10 cm s⁻¹ (52.0 ± 8.6 km), and lost 25% wet weight over that time. Vertical distribution differed between locations: modal depth was 2.5–5.0 and 10.0–12.5 m at leeward and windward sites, respectively. Over 80% of 71 larvae observed in situ had directional swimming trajectories. Larvae avoided NW bearings, with an overall mean SE swimming direction, regardless of the direction to nearest settlement habitat. Larvae made smaller changes between sequential bearings of swimming direction when swimming SE than in other directions, making it more likely they would continue to swim SE. When swimming NW, 62% of turns were left (more than in other directions), which would quickly result in swimming direction changing away from NW. This demonstrates the larvae knew the direction in which they were swimming and provides insight into how they achieved SE swimming direction. Although the cues used for orientation are unclear, some possibilities seemingly can be eliminated. Thus, A. curacao larvae near Lizard Island, on average swam into the average current at a speed equivalent to it, could do this for many hours, and chose different depths in different locations. These behaviours will strongly influence dispersal, and are similar to behaviour of other settlement-stage pomacentrid larvae that are older and larger.     

In situ ontogeny of behaviour in pelagic larvae of three temperate, marine, demersal fishes

The ontogeny of behaviour relevant to dispersal was studied in situ with reared pelagic larvae of three warm temperate, marine, demersal fishes: Argyrosomus japonicus (Sciaenidae), Acanthopagrus australis and Pagrus auratus (both Sparidae). Larvae of 5–14 mm SL were released in the sea, and their swimming speed, depth and direction were observed by divers. Behaviour differed among species, and to some extent, among locations. Swimming speed increased linearly at 0.4–2.0 cm s⁻¹ per mm size, depending on species. The sciaenid was slower than the sparids by 2–6 cm s⁻¹ at any size, but uniquely, it swam faster in a sheltered bay than in the ocean. Mean speeds were 4–10 body lengths s⁻¹. At settlement size, mean speed was 5–10 cm s⁻¹, and the best performing individuals swam up to twice the mean speed. In situ swimming speed was linearly correlated (R ²=0.72) with a laboratory measure of swimming speed (critical speed): the slope of the relationship was 0.32, but due to a non-zero intercept, overall, in situ speed was 25% of critical speed. Ontogenetic vertical migrations of several metres were found in all three species: the sciaenid and one sparid descended, whereas the other sparid ascended to the surface. Overall, 74–84% of individual larvae swam in a non-random way, and the frequency of directional individuals did not change ontogenetically. Indications of ontogenetic change in orientated swimming (i.e. the direction of non-random swimming) were found in all three species, with orientated swimming having developed in the sparids by about 8 mm. One sparid swam W (towards shore) when <10 mm, and changed direction towards NE (parallel to shore) when >10 mm. These results are consistent with limited in situ observations of settlement-stage wild larvae of the two sparids. In situ, larvae of these three species have swimming, depth determination and orientation behaviour sufficiently well developed to substantially influence dispersal trajectories for most of their pelagic period.     

Critical swimming speeds of late-stage coral reef fish larvae: variation within species,among species and between locations

The swimming abilities of larval fishes are important for their survival, potentially affecting their ability to avoid predators, obtain food and control dispersal patterns. Near settlement swimming abilities may also influence spatial and temporal patterns of recruitment. We examined Critical speed (U-crit) swimming ability in late stage larvae of 89 species of coral reef fishes from the Great Barrier Reef and the Caribbean. Coefficients of variation in U-crit calculated at the individual level were high (28.4%), and this was not explained by differences in size or condition factor of these same larvae. Among species U-crit ranged from 5.5 cm s⁻¹ to 100.8 cm s⁻¹ (mean=37.3 cm s⁻¹), with 95% of species able to swim faster than the average current speed around Lizard Island, suggesting that most species should be capable of influencing their spatial and temporal patterns of settlement. Inter-specific differences in swimming ability (at both the family and species levels) were significantly correlated with size and larval morphology. Correlations were found between swimming performance and propulsive area, fineness ratio and aspect ratio, and these morphological parameters may prove useful for predicting swimming ability in other taxa. Overall, the swimming speeds of larvae from the same families at the two locations were relatively similar, although the Lutjanidae and Acanthuridae from the Caribbean were significantly slower than those from the great barrier reef. Differences in swimming speed and body form among late stage larvae suggests that they will respond differently to factors influencing survival and transport during their pelagic phase, as well as habitat use following settlement.     

Effects of prey escape ability,flow speed,and predator feeding mode on zooplankton capture by barnacles

Experimental studies of feeding on zooplankton often involve the use of non-evasive Artemia spp. to represent zooplanktonic prey. Some zooplankton, however, such as copepods, are potentially evasive due to possession of effective predator-avoidance mechanisms such as high-speed escape swimming. In the present study, we compared the efficiencies with which non-evasive (A. salina) and evasive (copepods) zooplankton were captured by a sessile, suspension feeder, the coral-inhabiting barnacle Nobia grandis (Crustacea, Cirripedia). N. grandis specimens and zooplankton used in the present study were collected near Eilat, Israel in 1993. The effect of different flow speeds (from 0 to 14 cm s^-1) on captures of the two preys was also investigated. Additionally, we examined the effect of a flow-induced barnacle behavioral switch from active to passive suspension feeding, on zooplankton capture. Two video cameras were used to make close-up, three dimensional recordings of predator-prey encounters in a computer-controlled flow tank. Frame-by-frame video analysis revealed a highly significant difference (P< 0.001) in the efficiency with which A. salina and copepods were caught (A. salina being much more readily captured than copepods). After an encounter with cirri of feeding barnacles, copepods were usually able to swim out of the barnacles capture zone within one video frame (40 ms), by accelerating from a slow swimming speed (approximately 1.85 cm s^-1) to a mean escape swimming speed of 18.11 cm s^-1 (ca. 360 body lengths s^-1). This was not the case for A. salina nauplii, which usually remained in contact with cirri before being transferred to the mouth and ingested. Thus, experimental studies addressing the methodology of organisms feeding on zooplankton should consider that slow-swimming prey like Artemia sp. nauplii may only represent the non-evasive fraction of natural mesozooplankton assemblages.     

Riding Langmuir circulations and swimming in circles: a novel form of clustering behavior by the scyphomedusaLinuche unguiculata

Linuche unguiculata (Schwartz) seasonally forms patches in the Caribbean Sea and Indo-Pacific Ocean. Eighteen patches of medusae varying from about 500 m² to nearly 1 km² in area, were documented along the Belize barrier reef in March and April 1987, April 1988, and March and April 1990. The shape of each patch and the inter-medusa distances varied with wind velocity. At low wind speed (<4 m s^-1) patches were elliptical or circular and the individual medusae were separated by distances of 0.5 m, whereas at higher speeds windrows were evident and medusae were closer together. Windrows probably form by horizontal advection owing to convergence by Langmuir circulations. Because individual patches might exist for up to 4 mo as they drift downwind, and because winds of sufficient speed to produce Langmuir circulations do not always occur, a mechanism is necessary to maintain patch integrity during calms. In situ observations and in vitro video recording showed that the medusae swam in horizontal, near-surface, circular, clockwise trajectories. Although swimming speed was relatively high (up to 8 cm s^-1). net displacement velocity can be low (<1 cm s^-1). Thus, circular swimming probably reduces cluster breakup. Patch formation probably improves reproductive success by reducing sperm dilution.     

Aerobic metabolic rates of swimming juvenile mako sharks, <Emphasis Type="Italic">Isurus oxyrinchus</Emphasis>

The shortfin mako shark, Isurus oxyrinchus, is a highly streamlined epipelagic predator that has several anatomical and physiological specializations hypothesized to increase aerobic swimming performance. A large swim-tunnel respirometer was used to measure oxygen consumption (MO₂) in juvenile mako sharks (swimming under controlled temperature and flow conditions) to test the hypothesis that the mako shark has an elevated maintenance metabolism when compared to other sharks of similar size swimming at the same water temperature. Specimen collections were conducted off the coast of southern California, USA (32.94°N and 117.37°W) in 2001-2002 at sea-surface temperatures of 16.0–21.0°C. Swimming MO₂ and tail beat frequency (TBF) were measured for nine mako sharks [77–107 cm in total length (TL) and 4.4 to 9.5 kg body mass] at speeds from 28 to 54 cm s⁻¹ (0.27–0.65 TL s⁻¹) and water temperatures of 16.5–19.5°C. Standard metabolic rate (SMR) was estimated from the extrapolation to 0-velocity of the linear regression through the LogMO₂ and swimming speed data. The estimated LogSMR (±SE) for the pooled data was 2.0937 ± 0.058 or 124 mg O₂ kg⁻¹ h⁻¹. The routine metabolic rate (RMR) calculated from seventeen MO₂ measurements from all specimens, at all test speeds was (mean ± SE) 344 ± 22 mg O₂ kg⁻¹h⁻¹ at 0.44 ± 0.03 TL s⁻¹. The maximum metabolic rate (MMR) measured for any one shark in this study was 541 mg O₂ kg⁻¹h⁻¹ at 54 cm s⁻¹ (0.65 TL s⁻¹). The mean (±SE) TBF for 39 observations of steady swimming at all test speeds was 1.00 ± 0.01 Hz, which agrees with field observations of 1.03 ± 0.03 Hz in four undisturbed free-swimming mako sharks observed during the same time period. These findings suggest that the estimate of SMR for juvenile makos is comparable to that recorded for other similar-sized, ram-ventilating shark species (when corrected for differences in experimental temperature). However, the mako RMR and MMR are apparently among the highest measured for any shark species.     

Swimming behaviour of the saucer scallop Amusium balloti (Mollusca: Pectinidae)

The swimming performance of the saucer scallop Amusium balloti (Bernardi) was recorded from tests conducted in a natural environment in Shark Bay, Western Australia, in June, July and November 1984 and June and September 1985. Unlike all other scallops described in the literature, the swimming performance (both speed and distance) of A. balloti increases with size. Maximum distance swum in a single swimming event was 23.1 m, while the maximum cumulative distance swum (four swimming events) was 30.8 m. Swimming speeds for larger scallops were generally between 0.8 and 1.0 m s^-1 (1.6 and 2.0 knots), with a maximum speed of 1.6 m s^-1 (3.1 knots). Variations in swimming performance and response times with size and season are probably the major cause of variations in the scallop's vulnerability to fishing gear.     

Respiration rate and cost of swimming for Antarctic krill,<Emphasis Type="Italic"> Euphausia superba</Emphasis>, in large groups in the laboratory

Constructing realistic energy budgets for Antarctic krill, Euphausia superba, is hampered by the lack of data on the metabolic costs associated with swimming. In this study respiration rates and pleopod beating rates were measured at six current speeds. Pleopod beating rates increased linearly with current speed, reaching a maximum of 6 beats s^–1 at 17 cm s^–1. There was a concomitant linear increase in respiration rate, from 1.8 mg O₂ g_D^–1 h^–1 at 3 cm s^–1 to 8.0 mg O₂ g_D^–1 h^–1 at 17 cm s^–1. The size of the group tested (50, 100 and 300 krill) did not have a significant effect on pleopod beating rates or oxygen consumption (ANCOVA, F=0.264; P>0.05). The cost of transport reached a maximum of 75 J g^–1 km^–1 at 5 cm s^–1, and then decreased with increasing current speed to 29 J g^–1 km^–1. When considered in light of energy budgets for E. superba, these data indicate that the cost of swimming could account for up to 73% of total daily metabolic expenditure during early summer.Communicated by G.F. Humphrey, Sydney     

Tail beat frequency as a predictor of swimming speed and oxygen consumption of saithe (<Emphasis Type="Italic">Pollachius virens</Emphasis>) and whiting (<Emphasis Type="Italic">Merlangius merlangus</Emphasis>) during forced swimming

Oxygen consumption and tail beat frequency were measured on saithe (Pollachius virens) and whiting (Merlangius merlangus) during steady swimming. Oxygen consumption increased exponentially with swimming speed, and the relationship was described by a power function. The extrapolated standard metabolic rates (SMR) were similar for saithe and whiting, whereas the active metabolic rate (AMR) was twice as high for saithe. The higher AMR resulted in a higher scope for activity in accordance with the higher critical swimming speed (U _crit) achieved by saithe. The optimum swimming speed (U _opt) was 1.4 BL s⁻¹ for saithe and 1.0 BL s⁻¹ for whiting with a corresponding cost of transport (COT) of 0.14 and 0.15 J N⁻¹ m⁻¹. Tail beat frequency correlated strongly with swimming speed as well as with oxygen consumption. In contrast to swimming speed and oxygen consumption, measurement of tail beat frequency on individual free-ranging fish is relatively uncomplicated. Tail beat frequency may therefore serve as a predictor of swimming speed and oxygen consumption of saithe and whiting in the field.     

Energetics of underwater swimming in the great cormorant (Phalacrocorax carbo sinensis)

Resting metabolic rate (RMR), energy requirements and body core temperature were measured during underwater swimming in great cormorants (Phalacrocorax carbo sinensis) at the zoological garden in Neumünster, Germany, using gas respirometry and stomach temperature loggers. We used a 13 m long still water canal equipped with a respiration chamber at each end. Birds swam voluntarily in the canal at a mean speed of 1.51 ms^-1. Power input during underwater swimming averaged 31.4 W kg^-1. Minimal costs of transport of 19.1 J kg^-1 m^-1 were observed at a speed of 1.92 m s^-1. Body core temperature was stable in all birds within the first 60 min spent in the canal. After that, body temperature dropped at a rate of 0.14°C min^-1 until the birds voluntarily left the water. Our data indicate that great cormorants spend 2.7 times more energy than Adélie penguins (Pygoscelis adeliae) during underwater swimming. This can be essentially attributed to their poor insulation, their mode of locomotion underwater and differences in streamlining. RMR on land was related to body mass via VO₂=0.691 M^0.755 (where VO₂ is O₂-consumption in litre h^-1 and M is body mass in kg). In order to quantify the effects of external devices on energy consumption during underwater swimming, we tested a dummy data logger attached to the back of the cormorants as well as a ring on the leg. The ring had no apparent influence on the swimming energetics of the cormorants. In birds equipped with dummy loggers, swimming speed was not significantly influenced, but both power input and costs of transport increased by a mean of 19% for swimming speeds between 1.4 and 1.8 m s^-1.     

Locomotory activity and behaviour of the Antarctic teleostNotothenia coriiceps

The activity and behaviour of a free-living Antarctic fish,Notothenia coriiceps Richardson (formerlyN. neglecta), was investigated using a high-sensitivity, underwater TV camera at Signy Island, South Orkney Islands. Detailed observations of the 33 cm TL (total length) fish were made over a period of 6 d in austral summer (February 1992), for a total 69.5 h. Natural light at 2.5 m depth allowed viewing from 1 h before sunrise to 1 h after sunset. The fish stayed in a territory within 3 m of a small cave for >98% of the time, and made between 1 to 148 swims d^-1, of which 92.5% were brief (<15 s) feeding attempts. On average, 1.7% of each day was engaged in locomotion, including 1.2% swimming and 0.5% manoeuvring. Swimming was generally slow, at <2 body lengths s^-1, and labriform and subcarangiform modes were used alternately or in combination. Activity level (swims or displays per hour) was unaffected by tide, but was lower for 3 d when a wind speed >16 knots prevailed indicating that large waves reduced activity. A suspected diurnal activity rhythm was not statistically significant. The fish is an ambush-predator, and it took most of its prey from the water column but some off macroalgae or the seabed. Ventilation rate was slightly higher after activity, and peaked after an encounter with anotherN. coriiceps.     

Effect of temperature on the escape responses of larval herring,Clupea harengus

Herring (Clupea harengus L.) larvae from spring and autumn spawning stocks were reared at different constant temperatures from 5° to 17 °C. At equivalent developmental stages, the spring larvae were longer than the autumn larvae and the larvae reared at low temperatures were longer than those reared at high temperatures. At hatching and at the end of the yolk-sac stage, the larvae were induced, by a probe, to make C-start escape responses, which were recorded and analysed using a high-speed video recording at 400 frames s^-1. The response was rapid and of short duration. The tailbeat frequency and swimming speed were measured during the burst of swimming following the C-start at different test temperatures and in larvae with different temperature histories. The tail-beat frequency was strongly temperature-dependent, rising from 19 Hz at 5 °C to 37 Hz at 17 °C with no effect of temperature history, season or developmental stage. The burst-swimming speed ranged at hatching from 75 to 90 mm s^-1 at 5 °C to 110 to 160 mm s^-1 at 17 °C and at yolk resorption from 90–115 mm s^-1 at 5 °C to 175–190 mm s^-1 at 17 °C. The longer, spring-spawned larvae swam faster than the shorter autumn-spawned larvae. When the swimming speeds were expressed as body lengths (L) s^-1, these differences disappeared. Larvae swam from 7–9 L s^-1 at 5 °C to 15–20 L s^-1 at 17 °C at hatching, and from 8–9 L s^-1 at 5 °C to 15–17 L s^-1 at 17 °C at yolk resorption. There was, however, a significantly faster specific swimming speed by the larvae reared at 12 °C in spring 1991.Honorary Research Fellow of the Scottish Association for Marine ScienceUnfortunately, Karen Fretwell was drowned in an accident on 9 January 1993     

Feeding and swimming of lysianassid amphipods in a shallow cold-water bay

The potential for dispersal by lysianassid amphipods and their localization to carrion in a shallow cold-water bay in the Middle Saint Lawrence Estuary were assessed by means of endobenthic sampling, SCUBA observations, measures of swimming speeds, and by exposure of bait (50–100 g of fish) in traps. Seventy-five to 99.9% of animals attracted to traps were lysianassid amphipods belonging to five species. Lysianassid species were spatially segregated in the Bay at low tide but all were more or less dispersed at high tide. Second cohortAnonyx sarsi Steele and Brunel,Boeckosimus edwardsi andOnisimus littoralis (Krøyer) were more dispersed than the small first cohort individuals. Second cohortA. sarsi were crawlers or low (0–0.5 m off the bottom) suprabenthic swimmers in the day, but upper (0.5–2 m) suprabenthic swimmers at night. In contrast, first cohortA. sarsi were crawlers or low suprabenthic swimmers day-and-night, whileOrchomenella pinguis (Boeck) followed this swimming pattern at night but were generally akinetic in the day. Mean swimming speeds ofA. sarsi (13.6 cm s^-1) andOn. littoralis (12.1 cm s^-1) were 2 to 3 times greater than those ofOr. pinguis (7.4 cm s^-1) andPsammonyx nobilis (Stimpson) (4.4 cm s^-1). Catchability coefficients (i.e. ratio number of individuals per trap:endobenthic abundance) were 74 (A. sarsi), 8 (On. littoralis), 7 (Or. pinguis), and 0.7 (P. nobolis) m² of bottom. Gut content analysis indicated thatA. sarsi fed mostly on large carrion, whileOn. littoralis were markedly opportunistic, andOr. pinguis andP. nobilis relied on detritus, algae, and small crustaceans.     

Effects of current speed on filtration during suspension feeding in Oligometra serripinna (Echinodermata: Crinoidea)

Suspension feeding by the crinod Oligometra serripinna was studied at Lizard Island, Australia, in 1986. Video recordings were made of 90-m particles interacting with the filter of the crinoid in a laboratory flow chamber. A complete census of particles was possible because both the capture event and the filter area could be defined unequivocally. Also, because O. serripinna is a passive suspension feeder, a census of partcles could be made at different ambient current speeds without interference due to active pumping by the crinoid. Experiments were run at seven current speeds from 0.9 to 13.3 cm s^-1. Particles approaching the filter: (1) were captured, (2) passed through the filter without triggering a capture event, (3) passed through the filter after escaping from an unsuccessful capture event, or (4) were deflected around the filter. With increasing current speed, the proportion of deflections declined and the proportion of particles passing through rose: these results could be partially explained by the progressive widening of the spaces within the filter due to distortion of filter parts by the current. The proportion of captures (normalized to approaches) was comparatively low at 0.9 cm s^-1, rose to a relatively constant maximum from 1.7 to 6.4 cm s^-1, and then declined progressively at 9.5 and 13.3 cm s^-1. These proportions were translated into capture rates for whole crinoids by taking into consideration both the encounters with particles and the reduction of filter area by distortion of body parts at higher speeds. When plotted against current speed, capture rate peaked at 6.4 cm s^-1, which was close to the mean current speed that we measured on the reef in the microhabitat of O. serripinna.     

Determination of the swimming trajectory and speed of chain-forming dinoflagellate <Emphasis Type="Italic">Cochlodinium polykrikoides</Emphasis> with digital holographic particle tracking velocimetry

The marine dinoflagellate Cochlodinium polykrikoides is a harmful and highly motile algal species. To distinguish between the motility characteristics of solitary and chain-forming cells, the swimming trajectories and speeds of solitary cells and 2- to 8-cell chains of C. polykrikoides were measured using a digital holographic particle tracking velocimetry (PTV) technique. C. polykrikoides cells exhibited helical swimming trajectories similar to other dinoflagellate species. The swimming speed increased as the number of cells in the chain increased, from an average of 391 μm s⁻¹ (solitary cells) to 856 μm s⁻¹ (8-cell chain). The helix radius R and pitch P also increased as the number of cells in the chain increased. R increased from 9.24 μm (solitary cell) to 20.3 μm (8-cell chain) and P increased from 107 μm (solitary cell) to 164 μm (8-cell chain). The free thrust-generating motion of the transverse flagella and large drag reduction in the chain-forming cells seemed to increase the swimming speed compared to solitary cells. The measured swimming speeds agreed with those from field observations. The superior motility of chain-forming C. polykrikoides cells may be an important factor for its bloom, in addition to the factors reported previously.     

Acoustical identification of the concentration layer of a copepod species,Calanus euxinus

Swimming trajectories of Calanus euxinus Hulsemann in the Black Sea were studied using an echosounder at 120 and 200 kHz. C. euxinus were acoustically discriminated with respect to vertical migration and swimming speed, according to dissolved oxygen (DO) concentration and the timing of migrations. Species became torpid in water with DO values <0.5 mg l^у. The time spent swimming under DO conditions between 2 and 5 mg l^у was insignificant, and varied greatly from the 10% to 25% of total time spent swimming under normoxic conditions (5-10 mg l^у). C. euxinus formed a concentration layer in the water of 1-3 m thickness. Upward migration was completed in about 3.5 h, starting 2.5 h before and ending 1 h after sunset (average rate: 0.95 cm s^у) in summer. Species ascended discretely from the suboxic to the lower boundry of the cold intermediate layer (CIL) at 0.82 cm s^у, and passed up the CIL and thermocline fast (2.3 cm s^у). Downward migration took less time (2 h), starting ~1 h before and ending ~1 h after sunrise. Swimming speed within the thermocline and CIL was 2.7 cm s^у; copepods subsequently returned to daylight depth at a sinking speed of 0.57 cm s^у. Total time for C. euxinus to settle to their nocturnal depth layer was about 5 h.     

Temperature and plankton

The relationship between temperature and metabolism was studied in Artic copepods with regard to the concept of metabolic cold adaptation of polar poikilotherms. Temperature tolerance and respiration rates of the dominant copepods Calanus finmarchicus (Gunnerus), C. glacialis (Jaschnov), C. hyperboreus (Krøyer) and Metridia longa (Lubbeck), collected in Fram Strait, Greenland Sea, in July 1983, were studied at different temperatures. Temperature tolerance in the boreal C. finmarchicus was slightly higher than in the three Arctic species. Respiration rates at lower temperatures followed the Arrhenius equation in all species, with values for (temperature characteristics) between 11.05 and 22.95, corresponding to a Q₁₀ between 2.05 and 4.5. This increase in metabolic rate with rising temperature was not related to an increase of swimming activity, as was shown by videoanalysis. Activity was determined as average swimming speed and as frequency of certain locomotor patterns. Average swimming speed remained unchanged at all temperatures and was ca 1 cm s^-1 for all species, when only periods of active swimming were considered. The time spent with active swimming did not change with temperature in M. longa and C. finmarchicus, but decreased in c. glacialis. In C. hyperboreus it increased at 5°C and decreased again at higher temperatures. It is suggested that the increase in oxygen consumption is fully accounted for by the basal metabolism.     

The influence of swimming speed on sustained swimming performance of late-stage reef fish larvae

Replicate clutches of larvae were swum in a swimming flume at 4, 7, 10, 13, and 16 cm s^-1 and the time swum until exhaustion recorded. There was a significant relationship between sustained swimming time and swimming speed for both maximum (R²=0.77; P<0.05) and mean sustained swimming times (R²=0.78; P<0.05), with fish swimming at slower speeds swimming longer and covering greater total distances. The relationship observed agrees with theoretical principles relating increasing swimming speed with increasing drag. We used our data for Amphiprion melanopus, combined with published information, to predict the swimming speeds that other reef fish taxa should be able to maintain for significant lengths of time (12-48 h) using three different models. The results agree well with field estimates (R² values from 0.45 to 0.84), suggesting that there may be underlying factors influencing swimming ability in reef fish larvae that can be used to predict swimming abilities of different taxa. These models suggest that sustained swimming behaviour by reef fish larvae could have a much greater impact on modifying larval dispersal than previously thought.     

Life in a warm deep sea: routine activity and burst swimming performance of the shrimp<Emphasis Type="Italic"> Acanthephyra eximia</Emphasis> in the abyssal Mediterranean

Measurements of routine swimming speed, tail-flip escape responses, and oxygen consumptions were made of the deep-sea shrimp Acanthephyra eximia using autonomous landers in the Rhodos Basin at depths of up to 4,400 m and temperatures of 13–14.5°C. Routine swimming speeds at 4,200 m averaged 0.18 m s^–1 or 3.09 body lengths s^–1, approximately double those of functionally similar oceanic scavengers. During escape responses peak accelerations of 23 m s^–2 or 630.6 body lengths s^–2 were recorded, with animals reaching speeds of 1.61 m s^–1 or 34.8 body lengths s^–2. When compared to shallow-water decapods at similar temperatures these values are low for a lightly calcified shrimp such as A. eximia despite a maximum muscle mass specific power output of 90.0 W kg^–1. A preliminary oxygen consumption measurement indicated similar rates to those of oceanic crustacean scavengers and shallower-living Mediterranean crustaceans once size and temperature had been taken into account. These animals appear to have high routine swimming speeds but low burst muscle performances. This suite of traits can be accounted for by high competition for limited resources in the eastern Mediterranean, but low selective pressure for burst swimming due to reductions in predator pressure.Communicated by J.P. Thorpe, Port Erin     

Energy expenditure of swimming bottlenose dolphins (Tursiops truncatus)

The aim of our investigations was to determine, via oxygen and carbon-dioxide respirometry, how much energy dolphins (Tursiops truncatus) require when swimming at different speeds. Experiments were conducted on two female bottlenose dolphins (mean mass 162 kg) in the dolphinarium in Nuremberg Zoo, Germany, between March and August 1997. Animals were stationed in a respiration chamber for a minimum of 90 s after performing a variety of activities. We measured respiration frequency and oxygen requirements during (1) resting, (2) swimming at various velocities and (3) leaping to various heights. Resting metabolic rate of our bottlenose dolphins (2.15 W kg⁻¹) was comparable to previously published data. Metabolic rate in swimming dolphins increased to 2.47 W kg⁻¹ at 2 m s⁻¹, while leaps to 2.2 and 3 m height required a power input of 3.5 and 4 W kg⁻¹, respectively. Transport costs of swimming dolphins were lowest (1.16 J kg⁻¹ m⁻¹, corresponding to 0.12 J N⁻¹ m⁻¹) at a speed of 2.5 m s⁻¹, yielding an optimal range speed of between 1.9 and 3.2 m s⁻¹ (corresponding to minimum cost of transport ±10%). Breathing rates during all experiments correlated very well with oxygen consumption (r ² > 0.89) and could be used to derive metabolic rates in unencumbered dolphins at sea. Received: 18 December 1998 / Accepted: 27 April 1999