Settlement of <Emphasis Type="Italic">Macoma balthica</Emphasis> larvae in response to benthic diatom films

The role of multi-species benthic diatom films (BDF) in the settlement of late pediveliger larvae of the bivalve Macoma balthica was investigated in still-water bioassays and multiple choice flume experiments. Axenic diatom cultures that were isolated from a tidal mudflat inhabited by M. balthica were selected to develop BDF sediment treatments characterized by a different community structure, biomass, and amount of extracellular polymeric substances (EPS). Control sediments had no added diatoms. Although all larvae settled and initiated burrowing within the first minute after their addition in still water, regardless of treatment, only 48–52% had completely penetrated the high diatom biomass treatments after 5 min, while on average 80 and 69% of the larvae had settled and burrowed into the control sediments and BDF with a low diatom biomass (<3.5 μg Chl a g⁻¹ dry sediment), respectively. The percentage of larvae settling and burrowing into the sediment was negatively correlated with the concentration of Chl a and EPS of the BDF. This suggests higher physical resistance to bivalve penetration by the BDF with higher diatom biomass and more associated sugar and protein compounds. The larval settlement rate in annular flume experiments at flow velocities of 5 and 15 cm s⁻¹ was distinctly lower compared to the still-water assays. Only 4.6–5.8% of the larvae were recovered from BDF and control sediments after 3 h. Nonetheless, a clear settlement preference was observed for BDF in the flume experiments; i.e., larvae settled significantly more in BDF compared to control sediments irrespective of flow speed. Comparison with the settlement of polystyrene mimics and freeze-killed larvae led to the conclusion that active selection, active secondary dispersal and, at low flow velocities (5 cm s⁻¹), passive adhesion to the sediment are important mechanisms determining the settlement of M. balthica larvae in estuarine biofilms.     

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.     

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 swimming and settlement behaviour of larvae of an Indo-Pacific coral-reef fish, the coral trout Plectropomus leopardus (Pisces: Serranidae)

Late larvae of the serranid coral trout Plectropomus leopardus (Lacepède), captured in light traps, were released during the day both in open water and adjacent to two reefs, and their behaviour was observed by divers at Lizard Island, northern Great Barrier Reef. Coral trout larvae (n = 110) were present in light-trap catches from 18 November to 3 December 1997, including new moon (30 November). The swimming speed of larvae in open water or when swimming away from reefs was significantly greater (mean 17.9 cm s⁻¹) than the speed of larvae swimming towards or over reefs (mean 7.2 cm s⁻¹). Near reefs, larvae swam at average depths of 2.7 to 4.2 m, avoiding 0 to 2 m. In open water, swimming depth varied with location: larvae >1 km east of Lizard Island swam steeply downward to >20 m in 2 to 4 min; larvae >1 km west oscillated between 2.6 and 13 m; larvae 100 to 200 m east of Lizard Island oscillated between 0.8 and 15 m. Nearly all larvae swam directionally in open water and near reefs. In open water, the average swimming direction of all larvae was towards the island, and 80% (4 of 5) swam directionally (p < 0.05, Rayleigh's test). Larvae swam directionally over the reef while looking for settlement sites. The frequency of behaviours by larvae differed between two reefs of different exposure and morphology. Depending on site, 26 to 32% of larvae released adjacent to reefs swam to open water: of these, some initially swam towards or over the reef before swimming offshore. In some cases, offshore-swimming seemed to be due to the presence of predators, but usually no obvious cause was observed. Depending on the reef, 49 to 64% of the larvae settled. Non-predatory reef residents aggressively approached 19% of settlers. Between 5 and 17% of the larvae were eaten while approaching the reef or attempting to settle, primarily by lizardfishes but also by wrasses, groupers and snappers. A higher percentage of larvae settled in the second week of our study than in the first. Average time to settlement was short (138 s ± 33 SE), but some larvae took up to 15 min to settle. Average settlement depth was 7.5 to 9.9 m, and differed between locations. No settlement took place on reef flats or at depths <4.2 m. Larvae did not appear to be selective about settlement substrate, but settled most frequently on live and dead hard coral. Late-stage larvae of coral trout are capable swimmers with considerable control over speed, depth and direction. Habitat selection, avoidance of predators and settlement seem to rely on vision. Received: 7 July 1998 / Accepted: 26 January 1999     

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.     

Swimming ability of eels (<Emphasis Type="Italic">Anguilla rostrata,Conger oceanicus</Emphasis>) at estuarine ingress: contrasting patterns of cross-shelf transport?

The transport of eel early life stages may be critical to their population dynamics. This transport from ocean spawning to freshwater, estuarine and coastal nursery areas is a combination of physical and biological processes (including swimming behavior). In New Jersey, USA, the American eel (Anguilla rostrata) enters estuaries as glass eels (48.7–68.1 mm TL) in contrast to the Conger eel (Conger oceanicus) that enters as larger (metamorphosing) leptocephali (68.3–117.8 mm TL). To begin to understand the mechanisms of cross-shelf transport for these species, we measured the potential swimming capability (critical swimming speed, U _crit) under ambient conditions throughout the ingress season. A. rostrata glass eels were collected over many months (January–June) at a range of temperatures (4–21°C), with relative condition declining over the course of the ingress period as temperatures warmed. C. oceanicus occurred later in the season (April–June) and at warmer temperatures (14–24.5°C). Mean U _crit values for A. rostrata (11.7–13.3 cm s⁻¹) and C. oceanicus (14.7–18.6 cm s⁻¹) were comparable, but variable, with portions of the variability explained by water temperature, relative condition, ontogenetic stage, and fish length. Travel times to Little Egg Inlet, New Jersey, estimated using 50% U _crit values, indicate it would take A. rostrata ~30 and ~60 days to swim from the shelf edge and Gulf Stream, respectively. Travel times for C. oceanicus were shorter, ~20 days from the shelf edge, and ~45 days from the Gulf Stream. Despite differences in life stage, our results indicate both species are competent swimmers, and suggest they are capable of swimming from the Gulf Stream and/or edge of the continental shelf to estuarine inlets.     

Modification of light utilization for skeletal growth by water flow in the scleractinian coral <Emphasis Type="Italic">Galaxea fascicularis</Emphasis>

In this study, we tested the hypothesis that the importance of water flow for skeletal growth (rate) becomes higher with increasing irradiance levels (i.e. a synergistic effect) and that such effect is mediated by a water flow modulated effect on net photosynthesis. Four series of nine nubbins of G. fascicularis were grown at either high (600 μE m⁻² s⁻¹) or intermediate (300 μE m⁻² s⁻¹) irradiance in combination with either high (15–25 cm s⁻¹) or low (5–10 cm s⁻¹) flow. Growth was measured as buoyant weight and surface area. Photosynthetic rates were measured at each coral’s specific experimental irradiance and flow speed. Additionally, the instantaneous effect of water flow on net photosynthetic rate was determined in short-term incubations in a respirometric flowcell. A significant interaction was found between irradiance and water flow for the increase in buoyant weight, the increase in surface area, and specific skeletal growth rate, indicating that flow velocity becomes more important for coral growth with increasing irradiance levels. Enhancement of coral growth with increasing water flow can be explained by increased net photosynthetic rates. Additionally, the need for costly photo-protective mechanisms at low flow regimes could explain the differences in growth with flow.     

Effect of sub-lethal damage to juvenile colonies of massive Porites spp. under contrasting regimes of temperature and water flow

In this study, juvenile colonies of massive Porites spp. (a combination of P. lutea and P. lobata) from the lagoon of Moorea (W 149°50′, S 17°30′) were damaged and exposed to contrasting conditions of temperature and flow to evaluate how damage and abiotic conditions interact to affect growth, physiological performance, and recovery. The experiment was conducted in April and May 2008 and consisted of two treatments in which corals were either undamaged (controls) or damaged through gouging of tissue and skeleton in a discrete spot mimicking the effects of corallivorous fishes that utilize an excavating feeding mode. The two groups of corals were incubated for 10 days in microcosms that crossed levels of temperature (26.7 and 29.6°C) and flow (6 and 21 cm s⁻¹), and the response assessed as overall colony growth (change in weight), dark-adapted quantum yield of PSII (F _v/F _m), and healing of the gouged areas. The influence of damage on growth was affected by temperature, but not by flow. When averaged across flow treatments, damage promoted growth by 25% at 26.7°C, but caused a 25% inhibition at 29.6°C. The damage also affected F _v/F _m in a pattern that differed between flow speeds, with a 10% reduction at 6 cm s⁻¹, but a 4% increase at 21 cm s⁻¹. Regardless of damage, F _v/F _m at 21 cm s⁻¹ was 11% lower at 26.7°C than at 29.6°C, but was unaffected by temperature at 6 cm s⁻¹. The lesions declined in area at similar rates (4–5% day⁻¹) under all conditions, although the tissue within them regained a normal appearance most rapidly at 26.7°C and 6 cm s⁻¹. These findings show that the response of poritid corals to sub-lethal damage is dependent partly on abiotic conditions, and they are consistent with the hypothesis that following damage, calcification and photosynthesis can compete for metabolites necessary for repair, with the outcome affected by flow-mediated mass transfer. These results may shed light upon the ways in which poritid corals respond to biting by certain corallivorous fishes.     

Feeding behaviour of the sea cucumber <Emphasis Type="Italic">Cucumaria frondosa</Emphasis> (Echinodermata: Holothuroidea) in the laboratory and the field: relationships between tentacle insertion rate,flow speed,and ingestion

The sea cucumber, Cucumaria frondosa, is a benthic suspension feeder that captures food particles on its tentacles and then inserts them into its mouth one at a time. Previous studies have suggested that tentacle insertion rate (TIR) could be a useful indicator of food intake. The present study determined whether flow velocity affects TIR and whether TIR is a good indicator of ingestion. Video observations of sea cucumbers in Passamaquoddy Bay (45°01.70N, 66°55.74W) in August 1995 showed that TIRs increased with velocities up to 55 cm s⁻¹ and decreased steadily at flows above that up to 130 cm s⁻¹. In October 2006, laboratory flume studies were carried out on specimens collected from the same site in the previous August. Temperature and salinity (12°C and 32) in the flume were the same as in the field at the time of collection. There was high individual variation in feeding behavior at free-stream velocities of 4–40 cm s⁻¹ and TIR was independent of flow. As the number of tentacle insertions increased in the flume experiments, the amount of chloropigments in the digestive tracts of the sea cucumbers also increased. This suggests that TIR, which can be measured non-intrusively using remote video techniques, could be a good indicator of feeding behavior and ingestion in C. frondosa.     

Automated method for the determination of total arsenic and selenium in natural and drinking water by HG-AAS

A multicommutated flow system was designed and evaluated for the determination of total arsenic and selenium by Hydride Generation Atomic Absorption Spectrometry (HG-AAS). It was applied to the determination of arsenic and selenium in samples of natural and drinking water. Detection limits were 0.46 and 0.08 μg l⁻¹ for arsenic and selenium, respectively; sampling frequency was 120 samples h⁻¹ for arsenic and 160 samples h⁻¹ for selenium. Linear ranges found were 1.54–10 μg l⁻¹ (R = 0.999) for arsenic and 0.27–27 μg l⁻¹ (R = 0.999) for selenium. Accuracy was evaluated by spiking various water samples and using a reference material. Recoveries were in the range 95–116%. Analytical precision (s _r (%), n = 10) was 6% for both elements. Compared with the Standard Methods, APHA, 3114B manual method, the system consumes at least 10 times less sample per determination, and the quantities of acid and reducing agent used are significantly lower with a reduction in the generation of pollutants and waste. As an additional advantage, the system is very fast, efficient and environmentally friendly for monitoring total arsenic and selenium levels in waters.     

Physiological effects of the detergent linear alkylbenzene sulphonate on blue mussel larvae (Mytilus edulis) in laboratory and mesocosm experiments

A series of laboratory (short-term exposure in small beakers) studies and a 19 d mesocosm (6 m³ polyethylene bags filled with fjord water) study were conducted on blue mussel, Mytilus edulis, larvae and plantigrades exposed to a concentration gradient of the detergent linear alkylbenzene sulphonate (LAS, 0 to 39 mg l⁻¹). LAS is increasingly found in nearshore environments receiving wastewater from urban treatment plants. The aims were to observe physiological effects on swimming, grazing and growth in the laboratory and effects on settling and population development at in situ conditions (in field mesocosms) in order to evaluate the damages on ciliated meroplankton caused by LAS. In the laboratory the larvae showed a 50% mortality at 3.8 mg LAS l⁻¹ after 96 h exposure whether or not food was provided. Additionally the swimming behaviour was affected at 0.8 mg LAS l⁻¹ (i.e. a more compact swimming track, a smaller diameter of the swimming tracks, and reduced swimming speed). The larval particle grazing was reduced 50% at 1.4 mg LAS l⁻¹. The specific growth rate of the larvae was reduced to half at 0.82 mg LAS l⁻¹ over 9 d. During the mesocosm experiment, the larval population showed a dramatic decrease in abundance within 2 d at concentrations as low as 0.08 mg LAS l⁻¹, both due to a significantly increased mortality, but also due to settling. The settling success was reduced at the same LAS concentration as that at which mortality was observed to increase significantly. In addition to reduced settling rate, the larvae showed delayed metamorphosis and reduced shell growth as a response to LAS. Our hypothesis that the larval ciliary apparatus, crucial for normal swimming, orientation, and settling behaviours and for particle uptake, was damaged due to LAS exposure is supported by our results. This is confirmed by the physiological data (grazing, growth) and in the direct video-based observations of larval performance (swimming) and provides a reasonable explanation for what was observed in the bags (abundance, settling, mortality). These physiological effects on blue mussel larvae/plantigrades occurred at LAS concentrations reported to occur in estuarine waters. Received: 15 January 1997 / Accepted: 12 February 1997     

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.     

Experimental investigation into the effects of suspended sediment on fertilisation, larval survival and settlement in a scleractinian coral

Laboratory and field experiments were used to determine whether high (≃100 mg l⁻¹), low (≃50 mg l⁻¹) and control (≃0 mg l⁻¹) levels of suspended sediment affected fertilisation, larval survival, and larval settlement in the scleractinian coral Acropora digitifera (Dana, 1846). Both high- and low-sediment treatments significantly decreased fertilisation, but post-fertilisation embryonic development was not inhibited by suspended sediments. Larval survival and larval settlement were significantly reduced in high- and low-sediment treatments. No difference was found between high- and low-sediment treatments in any of the three post-spawning processes investigated, suggesting that they are susceptible to sediment concentrations which are not exceptionally high even under natural conditions (>50 mg l⁻¹). The introduction of an additional stress in the form of high levels of suspended sediments coupled with naturally high variability in recruitment may have a considerable effect on the successful supply and settlement of coral larvae to a reef. Given that many coral communities are open reproductive systems, the consequences of disturbance events are not likely to be restricted to the impact area. Recruitment to a population may be reduced significantly in the presence of high levels of suspended sediments because of effects on larval survival and settlement. Recruitment of larvae to adjacent populations may also be affected due to a decreased fertilisation success and potential increases in mortality of larvae passing through the affected site. Received: 13 August 1998 / Accepted: 22 July 1999     

Prolonged exposure to low dissolved oxygen affects early development and swimming behaviour in the gastropod Nassarius festivus (Nassariidae)

Effects of low dissolved oxygen on early development and swimming behaviour of veliger larvae of the scavenging gastropod Nassarius festivus were studied. Embryonic development was significantly delayed when dissolved oxygen level was reduced to 3.0 mg O₂ l⁻¹ and no embryo hatched successfully at 0.5 mg O₂ l⁻¹. Veliger larvae hatched at 4.5 mg O₂ l⁻¹ had significantly smaller velar lobe, shell length and shell width. Median 48-h LC₅₀ value of the veliger larvae was estimated at 1.25 mg O₂ l⁻¹ with lower swimming speed (swimming velocity and dispersal velocity) being recorded for the survivors exposed to reduced oxygen levels. The percentage of veliger larvae that developed into crawling juveniles was significantly reduced and metamorphosis was delayed at 4.5 mg O₂ l⁻¹ whereas all larvae at 3.5 mg O₂ l⁻¹ died before they underwent metamorphosis. Juveniles developed at 4.5 mg O₂ l⁻¹ were also smaller than those at 6.0 mg O₂ l⁻¹. Results indicated that dissolved oxygen levels well above hypoxia levels (2.8 mg O₂ l⁻¹) have already had significant impact on the hatching success and larval development in gastropods, which may lead to long-term decreases in population growth.     

Foraging behaviour of larval cod (<Emphasis Type="Italic">Gadus morhua</Emphasis>) at low light intensities

The ability to forage at low light intensities can be of great importance for the survival of fish larvae in a pelagic environment. Three-dimensional silhouette imaging was used to observe larval cod foraging and swimming behaviour at three light intensities (dusk ~1.36 × 10⁻³ W/m², night ~1.38 × 10⁻⁴ W/m² and darkness ~3.67 × 10⁻⁶ W/m²) at 4 different ages from 6 to 53 days post-hatch (dph). At 6 dph, active pursuit of prey was only observed under dusk conditions. Attacks, and frequent orientations, were observed from 26 dph under night conditions. This was consistent with swimming behaviour which suggested that turn angles were the same under dusk and night conditions, but lower in darkness. Cod at 53 dph attacked prey in darkness and turn angles were not different from those under other light conditions. This suggests that larvae are still able to feed at light intensities of 3.67 × 10⁻⁶ W/m². We conclude that larval cod can maintain foraging behaviour under light intensities that correspond to night-time at depths at which they are observed in the field, at least if they encounter high-density patches of prey such as those that they would encounter at thin layers or fronts.     

Halogenated metabolites in two marine polychaetes and their planktotrophic and lecithotrophic larvae

This study investigated the occurrence and ontogenetic changes of halogenated secondary metabolites in planktotrophic and lecithotrophic larvae and adults of two common, infaunal polychaetes, Streblospio benedicti (Spionidae) and Capitella sp. I (Capitellidae), with different life-history traits. S. benedicti contains at least 11 chlorinated and brominated hydrocarbons (alkyl halides) while Capitella sp. I contains 3 brominated aromatic compounds. These halogenated metabolites are potential defense compounds benefiting both larvae and adults. We hypothesized that: (1) planktotrophic larvae contain halogenated metabolites because they are not protected by adult defenses, (2) quantitative and qualitative variation of planktotrophic larval halogenated metabolites parallels that of adults, and (3) brooded lecithotrophic larvae initiate the production of halogenated metabolites only after metamorphosis. To address these hypotheses, volatile halogenated compounds from polychaete extracts were separated using capillary gas chromatography and identified and quantified using mass spectrometry with selected ion monitoring. All four life stages (pre- and post-release larvae, new recruits, adults) of both S. benedicti and Capitella sp. I contained the halogenated metabolites previously identified from adults. This is the first report of halocompounds identified and quantified in polychaete larvae. Allocation of potential defense compounds to offspring varied as a function of species, feeding type and developmental stage. Pre-release larvae of S. benedicti with planktotrophic development contained the lowest concentration of total halogenated metabolites (1.75 ± 0.65 ng mm⁻³), post-release and new recruits contained intermediate concentrations (8.29 ± 1.72 and 4.73 ± 2.63 ng mm⁻³, respectively), and planktotrophic adults contained significantly greater amounts (28.9 ± 9.7 ng mm⁻³). This pattern of increasing concentrations with increasing stage of development suggests synthesis of metabolites during development. Lecithotrophic S. benedicti post-release larvae contained the greatest concentrations of halometabolites (71.1 ± 10.6 ng mm⁻³) of all S. benedicti life stages and developmental types examined, while the amount was significantly lower in new recruits (34.0 ± 15.4 ng mm⁻³). This pattern is consistent with a previously proposed hypothesis suggesting a strategy of reducing potential autotoxicity during developmental transitions. Pre-release lecithotrophic larvae of Capitella sp. I contained the highest concentration of total halogenated metabolites (1150 ± 681 ng mm⁻³), whereas the adults contained significantly lower total amounts (126 ± 68 ng mm⁻³). All concentrations of these haloaromatics are above those known to deter predation in previously conducted laboratory and field trials. As a means of conferring higher larval survivorship, lecithotrophic females of both species examined may be expending more energy on chemical defenses than their planktotrophic counterparts by supplying their lecithotrophic embryos with more of these compounds, their precursors, or with energy for their synthesis. This strategy appears common among marine lecithotrophic larval forms. Received: 14 July 1999 / Accepted: 20 January 2000     

Macroalgal morphology mediates particle capture by the corallimorpharian <Emphasis Type="Italic">Corynactis californica</Emphasis>

The shallow kelp forest at Santa Catalina Island, California (33.45 N, −118.49 W) is distinguished by several canopy guilds ranging from a floating canopy (Macrocystis pyrifera), to a stipitate, erect understory canopy (Eisenia arborea), to a short prostrate canopy just above the substratum (Dictyopteris, Gelidium, Laminaria, Plocamium spp.), followed by algal turfs and encrusting coralline algae. The prostrate macroalgae found beneath E. arborea canopies are primarily branching red algae, while those in open habitats are foliose brown algae. Densities of Corynactis californica, are significantly greater under E. arborea canopies than outside (approximately 1,200 versus 300 polyps m⁻² respectively). Morphological differences in macroalgae between these habitats may affect the rate of C. californica particle capture and serve as a mechanism for determining polyp distribution and abundance. Laboratory experiments in a unidirectional flume under low (9.5 cm s⁻¹) and high (21 cm s⁻¹) flow speeds examined the effect of two morphologically distinct macroalgae on the capture rate of Artemia sp. cysts by C. californica polyps. These experiments (January–March 2006) tested the hypothesis that a foliose macroalga, D. undulata, would inhibit particle capture more than a branching alga, G. robustum. G. robustum, found predominantly under the E. arborea canopy did not affect particle capture. However, D. undulata, found predominantly outside of the canopy, inhibited particle capture rates by 40% by redirecting particles around C. californica polyps and causing contraction of the feeding tentacles. These results suggest that the morphology of flexible marine organisms may affect the distribution and abundance of adjacent passive suspension feeders.     

Development of the nasal olfactory organs in the larvae, settlement-stages and some adults of 14 species of Caribbean reef fishes (Labridae, Scaridae, Pomacentridae)

Larval fishes likely use a variety of settlement cues to locate and navigate toward the habitats they will inhabit as juveniles. Information about the morphology and state of development of the sensory organs of larval stages of fishes provides insight into their capabilities at the time of settlement. The peripheral olfactory organ of wild-caught late-stage larvae and early juveniles and some adults of 14 species of the Caribbean reef fishes wrasses (Labridae), parrot fishes (Scaridae) and damselfish (Pomacentridae) were examined using scanning electron microscopy and compared in terms of settlement specificity. Ages in days after hatching and days post-settlement were determined from the otoliths. Morphology of the nares and the olfactory epithelium are described for these species by stage. The separation of the anterior and posterior nares occurred before settlement in the labrids but in some specimens of scarids this separation was not complete by the time of settlement. Densities of ciliated and microvillous receptor cells and non-sensory ciliated epithelial cells were calculated. Densities of ciliated receptor cells ranged from 0.389 μm⁻² in a specimen of Thallasoma bifasciatum to 0.0057 μm⁻² in Bodianus rufus and of microvillous receptor cells from 0.038 μm⁻² in a Clepticus parrae juvenile to 0.266 μm⁻² in a juvenile Doratonotus megalepis. Densities of non-sensory cilia, also associated with high olfactory ability, were also high. The olfactory organ in wrasses is well developed prior to settlement and is comparable to that of adult fishes. The possible role of olfaction in larval schooling, reef cue detection and orientation toward habitat at settlement is discussed.

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Localisation of biogenic amines in larvae of Bugula neritina (Bryozoa: Cheilostomatida) and their effects on settlement

Distributions of serotonin and catecholamines in larvae of the marine bryozoan Bugula neritina (Bryozoa: Cheilostomatida) were investigated using immunohistochemistry with anti-serotonin antiserum and glyoxylic acid–induced fluorescence histochemistry. Anti-serotonin immunoreactive substances and glyoxylic acid–induced fluorescent substances had similar distributions in the equatorial neuromuscular ring, the neural plexus, the paired axial neuromuscular cords, and tracts connecting the neural plexus to ciliated cells bordering the pyriform organ. The effects of dopamine, noradrenaline, adrenaline, tyramine, octopamine, synephrine and serotonin, at 10⁻⁴, 10⁻⁵ and 10⁻⁶  M, on settlement were analysed. In filtered seawater, 98% of larvae settled in 3 h, but only 11%, 3% and 6% total settlement was observed after 8 h in 10⁻⁴  M dopamine, 10⁻⁴  M serotonin and 10⁻⁵  M serotonin, respectively. Total settlement was 70% in 10⁻⁴  M noradrenaline, 80% in 10⁻⁴  M adrenaline and 60% in 10⁻⁴  M tyramine. Less than 60% settlement was observed in 10⁻⁴ and 10⁻⁵  M octopamine and synephrine. Serotonin's inhibitory effect on settlement was mimicked by a range of serotonin receptor agonists and antagonists, among which 5-carboxamidotryptamine was the most potent. Received: 19 March 1999 / Accepted: 11 October 1999     

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.