Energy demand of larval and juvenile Cape horse mackerels, Trachurus capensis, and indications of hypoxia tolerance as benefit in a changing environment

Trachurus capensis is an important fisheries resource in the degraded Namibian upwelling ecosystem. Food supply and shoaling of hypoxic zones are hypothesised to influence the species’ recruitment success. This paper is the first to quantify energy requirements and hypoxia tolerance of larval and juvenile stages of a Trachurus species. We measured normoxic respiration rates of T. capensis with a size range from 0.001 to 20.8 g wet mass (WM) collected off Cape Town (33.9°S, 18.5°E, 12/2009) and in the northern Benguela (17–24°S, 11–15°E, 02/2011). Routine metabolic rate (RMR) and standard routine rate (SRR) (mg O₂ h^?1) followed the allometric functions RMR = 0.418 WM^0.774 and SRR = 0.275 WM^0.855, respectively. Larvae and juveniles had comparatively high metabolic rates, and the energy demand of juveniles at the upper end of the size range appeared too high to be fuelled by a copepod diet alone. T. capensis’ early life stages showed a high tolerance to hypoxic conditions. RMR in larvae did not change until 30 % O₂sat at 18 °C. In juveniles, critical oxygen saturation levels were low (P_C for SRR = 11.2 ± 1.7 % O₂sat and P_C for RMR = 13.2 ± 1.6 % O₂sat at 20 °C) and oxy-regulation effective (regulation index = 0.78 ± 0.09). A high hypoxia tolerance may facilitate the retention of larvae in near-shore waters providing favourable feeding conditions and allowing juveniles to exploit food resources in the oxygen minimum zone. These mechanisms seem to well adapt T. capensis to a habitat affected by spreading hypoxic zones and probably enhance its recruitment success.     

Thermal tolerance of early development in tropical and temperate sea urchins: inferences for the tropicalization of eastern Australia

The thermal envelope of development to the larval stage of two echinoids from eastern Australia was characterized to determine whether they fill their potential latitudinal ranges as indicated by tolerance limits. The tropical sand dollar, Arachnoides placenta, a species that is not known to have shifted its range, was investigated in Townsville, northern Australia (19°20′S, 146°77′E), during its autumn spawning season (May 2012). The subtropical/temperate sea urchin, Centrostephanus rodgersii, a species that has undergone poleward range expansion, was investigated in Sydney, southern Australia (33°58′S, 151°14′E), during its winter spawning season (August 2012). The thermal tolerance of development was determined in embryos and larvae reared at twelve temperatures. For A. placenta, the ambient water temperature near Townsville and experimental control were 24 °C and treatments ranged from 14 to 37 °C. For C. rodgersii, ambient Sydney water temperature and experimental control were 17 °C, and the treatment range was 9–31 °C. A. placenta had a broader developmental thermal envelope (14 °C range 17–31 °C) than C. rodgersii (9 °C range 13–22 °C). Both species developed successfully at temperatures well below ambient, suggesting that cooler water is not a barrier to poleward migration for either species. Both species presently live near the upper thermal limits for larval development, and future ocean warming could lead to contractions of their northern range limits. This study provides insights into the factors influencing the realized and potential distribution of planktonic life stages and changes to adult distribution in response to global change.     

Identification of archaeogastropod larvae from a hydrothermal vent community

Dispersal is essential in order that endemic species living in ephemeral, patchy vent environments may persist over evolutionary time. Quantitative field studies of larval dispersal, however, require specieslevel identification of the larval forms because each individual must be distinguished from related vent species, and from non-vent species living in the surrounding deep-sea environment. Methods for culturing these larvae to an identifiable stage have not yet been developed. To solve the larval identification problem for the archaeogastropod molluscs (a prominent component of vent communities), we used a scanning electron microscope (SEM) to image shells of larvae collected in the water column near vents along the East Pacific Rise (9°40′ to 9°50′N; 104°W). Larval shell size, shape and ornamentation were compared to protoconchs retained in juvenile or adult shells of identified species, and used to assign five larval groups unequivocally to species (Cyathermia naticoides Warén and Bouchet, 1989; Neomphalus fretterae McLean, 1981; Clypeosectus delectus McLean, 1989; Rhynchopelta concentrica McLean, 1989; and Lirapex granularis Warén and Bouchet, 1989) and seven groups tentatively to species or genus [Lepetodrilus spp. (three groups); Gorgoleptis sp; Peltospira ?operculata McLean, 1989; and ?Melanodrymia sp. (two groups)].     

Physiological development of brooded larvae from two pocilloporid corals in Taiwan

In southern Taiwan, brooded larvae of Pocillopora damicornis and Seriatopora caliendrum are released year-round in synchrony with new moons, and each larval release occurs over multiple days. Using P. damicornis and S. caliendrum as a model system, we describe within-brood variation in larval phenotypes and test for release-day effects that influence larval performance in the pelagic phase. Research was conducted in 2010 using larvae from corals collected in June and July from Nanwan Bay (21°56.179??N, 120°44.85??E). In June, larval phenotypes of both species were characterized immediately following release, and their competency to settle assessed. In July, larvae of P. damicornis were collected on 3?days over the peak release period and incubated for 7?C11?days at 28.0?°C and 320???mol?quanta?m^?2?s^?1; their phenotypes and settlement competency were measured every 2?days. P. damicornis larvae released close to peak release were 1.6 times larger in size, contained twice the number of Symbiodinium larva^?1, and were 44?% more likely to settle in the first 24?h than larvae released early in the brood. In addition, peak-release larvae respired at a lower rate than larvae released late in the brood. Similarly, S. caliendrum larvae released close to peak release were 1.4 times larger in size and were 33?% more likely to settle in the first 5?h than larvae released early in the brood. In July, P. damicornis larvae differed between early (2?days prior to peak), peak, and late (2?days after peak) release. Protein content of early-release larvae was lower than peak- and late-release larvae, and this difference persisted throughout the development. Further, release day affected the way larval respiration varied throughout development. By showing that brooded coral larvae differ between release days and display maternal effects influencing performance in the swimming phase, our results suggest that pocilloporid corals utilize bet-hedging to increase reproductive success.     

Genetic connectivity of the ecosystem engineer Perumytilus purpuratus north to the 32°S southeast Pacific ecological discontinuity

Connectivity in benthic marine animals with complex life cycles occurs primarily during the pelagic larval stage and depends deterministically on oceanographic dynamics. The scale of such larval dispersal is highly uncertain due the difficulty of direct measurement and poor knowledge of larval dynamics and ocean flow variability. This study characterizes the pattern of genetic connectivity in the ecosystem engineer Perumytilus purpuratus between latitudes 23°S and 33°S, which includes the ecological discontinuity reported for many taxa north to 32°S at the southeast Pacific. The genetic discontinuity observed in P. purpuratus around 26°S is described herein while that detected at 28°S is in line with the ecological discontinuity (in coverage, recruitment and density) previously reported for this mussel between 28°S and 32°S. Both discontinuities delimitate two major gene pools upon Bayesian inferences on geographical variation of five microsatellite loci. Interestingly, marker Pepu1 was responsible for most variation between pools and was potentially under selection. In fact, inferences excluding Pepu1 produced a single gene pool (k = 1) in central-northern Chile. The IBD connectivity pattern observed among P. purpuratus beds distributed in the interval 23°S–33°S is congruent with processes driven by larval dynamics, and the dominant equatorward Humboldt Current along a coast largely unaffected by ice during the last Pleistocene glaciation. However, the selective scenario unveiled by microsatellite Pepu1 inside the 25°S–28°S ecological discontinuity is consistent with selective processes associated with specific mesoscale properties operating in this area. This study highlights the usefulness of integrating different oceanographic scales, ecological data and population genetics to better understand connectivity of benthic marine species.     

Where are larvae of the vermetid gastropod <Emphasis Type="Italic">Dendropoma maximum</Emphasis> on the continuum of larval nutritional strategies?

This study evaluated whether larvae of the Indo-Pacific vermetid gastropod Dendropoma maximum are obligate planktotrophs, or whether they exhibit an intermediate feeding strategy. Experiments were conducted in Moorea, French Polynesia (149°50′W, 17°30′S), Sep–Oct 2009, to examine D. maximum larval growth and metamorphic responses to different diets and amounts of food. Dendropoma maximum larvae required particulate food to undergo metamorphosis, but were able to survive and grow in the absence of food for up to 20 days. Larvae in Low and Unfed food treatments exhibited phenotypic plasticity by growing a larger velum (the larval feeding structure) compared with those in high food. Unfed D. maximum larvae had a slower initial growth rate; however, by 11-day post-hatch fed and unfed larvae had converged on the same mean shell height (553 μm), which was only 10% larger than the initial size at hatching. Therefore, although the nutritional strategy of D. maximum larvae is best described as obligate planktotrophy, it appears to approach an intermediate feeding strategy.     

Spatial patterns of ichthyoplankton distribution in relation to hydrographic features in the Northern Benguela region

The spatial distribution of the most abundant eggs and larvae of teleost fish species on the continental shelf and slope off the northern Benguela region was studied in April 1986. The horizontal and vertical distribution of eggs and larvae were analysed together with environmental data, in order to determine patterns of ichthyoplankton distribution. Both species composition and relative egg and larval abundance levels exhibited important latitudinal differences during a period of quiescent upwelling with an intense intrusion of Angolan water into the system. Larval diversity was higher in the northern part of the study area, where, because of the intrusion of the warmer Angolan water, the water column was more stratified than in the southern part, where the affect of upwelling of South Atlantic Central Water was continuous and only a few species spawned. The frontal zone appeared to be a nursery ground for the most important pelagic species of the region:Trachurus trachurus capensis, Engraulis capensis, andSardinops ocellatus. Vertical egg and larval distributions showed evidence of stratification, with highest concentrations located in the uppermost 50 m. In comparison, during periods of intense upwelling, longitudinal gradients were responsible for the horizontal distribution of ichthyoplankton, and the vertical distribution of eggs and larvae were much more extensive because of the greater mixing of the water column.     

Effects of self-fertilisation and other factors on the early development of the scallop Pecten maximus

Adult Pecten maximus (L.) were dredged off north-east Anglesey, Wales, UK, during 1981. A 2×5 factorial mating was carried out involving self- and cross-fertilisation and the use of stripped spermatozoa. Assessments of yield, normality, and larval size were made at the D larva stage and larval size and mortality were measured after a fortnight's growth of the veligers. Underlying genetic variation was evident at all stages, with egg and sperm generally having a significant interactive effect. Cultures sired with stripped spermatozoa had fewer larvae, with more abnormality, grew slower and suffered higher mortality than most other cultures. Larvae from selfed cultures grew significantly slower than all other larvae. Data from past larval cultures also show that selfed larvae have a reduced growth rate. it is suggested that stripped spermatozoa may interfere with egg and sperm interaction at fertilisation, thus reducing the viability of larvae. On the other hand, the poor growth rate of selfed larvae is probably due to overall reduced heterozygosity compared to outbred larvae.     

Some observations of factors affecting embryonic and larval growth of Pollicipes polymerus (Cirripedia: Lepadomorpha) in vitro

Observations on the effects of several environmental conditions in embryonic and larval cultures are reported for the gooseneck barnacle Pollicipes polymerus (Sowerby, 1833). When the growth rates of control embryos kept in finger bowls at ambient-temperature sea water (ca. 12°–15°C) were compared to those of embryos grown under variously modified circumstances, it was found that growth rates were faster under conditions closely simulating the adult barnacle mantle-cavity (e.g. darkness and aeration). Addition of antimicrobial drugs and reduction of egg-mass size also promoted fast growth and development. Nauplii were fed several species of algae; only three species promoted growth to Stage V or older. Naupliar growth was fastest in larvae fed the combination of Prorocentrum micans/Platymonas sp. Larvae fed Prococentrum micans/pennate diatom grew slowest, and most larvae died at Naupliar Stage III. Interactions between algal species may have affected their nutritional value for Pollicipes polymerus nauplii. Less larval activity, slower growth rates, and higher mortality rates were observed in individuals given small amounts of food. The development of embryos and larvae in culture depends greatly on the culture conditions. These conditions should be described if comparison of timetables and envents are to be made between studies.     

Comparative larval development of the shipworms Bankia gouldi and Teredo navalis

Larvae of Bankia gouldi (Bartsch) and Teredo navalis L. were reared in the laboratory at various temperatures and salinities. T. navalis spawned at lower temperatures than B. gouldi. T. navalis larvae were released at temperatures from 13° to 30°C; in B. gouldi, spawning occurred from 17.5° to 30°C. Both species released offspring at salinities of 20 and 30‰. Larvae of the two species can be distinguished morphometrically at the earliest pelagic veliger stage and in the pediveliger stage. Average dimensions of newly released T. navalis larvae are 88 x 75 μ (length x height), while the youngest B. gouldi veligers measure 61 x 50 μ. Pediveligers of T. navalis (205 x 239 μ) are smaller than those of B. gouldi (221 x 260 μ). At other stages of larval development the two species appear so similar that they cannot yet readily be distinguished. Under laboratory conditions of 25°C and 30 ‰, the free-swimming life of B. gouldi to the pediveliger stage was about 10 days longer than that of T. navalis (25 and 15 days, respectively). Incubated larval development of T. navalis was estimated to be 5 days at 25°C. Potential competition between larval stages of the two species, and modification of settling behavior by dissolved humic material (Gelbstoff), is discussed.     

Substratum preferences in planula larvae of two species of scleractinian corals, Goniastrea retiformis and Stylaraea punctata

To test whether coral planulae recruit randomly to different coral reef habitats or have specific substratum preferences, the settling behavior of planulae from two shallow water coral species from Pago Bay, Guam (13°25.02N, 144°47.30E) were examined in the laboratory in June and July of 1995. Goniastrea retiformis is generally restricted to the shallow reef front (<10 m depth) in areas dominated by crustose coralline algae (CCA), while Stylaraea punctata is abundant on inner reef flats were CCA coverage is low and sand and carbonate rubble covered by biofilms is common. When presented with four substrata (1) carbonate rock scrubbed free of biofilm and dried as a control, (2) the CCA Hydrolithon reinboldii, (3) the CCA Peyssonelia sp., and (4) naturally conditioned carbonate rubble covered by a biofilm, G. retiformis larvae showed a significant preference for H. reinboldii, and S. punctata larvae for the carbonate biofilm treatment. The preference shown by S. punctata larvae for biofilmed surfaces did not diminish with increasing larval age up to 11 days. These results suggest that the larvae of both species are capable of habitat selection, and that the preferred substrata among those tested bears a relationship to the habitats in which adult colonies were found. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Larvae of benthic invertebrates in hydrothermal vent plumes over Juan de Fuca Ridge

Larvae of benthic invertebrates collected in the water column above Juan de Fuca Ridge show distinct variations in abundance and composition in, and away from, the neutrally-buoyant hydrothermal plume emanating from underlying vents. Larvae of vent gastropods (Lepetodrilus sp. and two peltospirid species) occur in significantly higher abundances in the plume than away from it (mean abundance=21.0 individuals 1000 m^?3 vs 1.4 individuals 1000 m^?3), and larvae of vent bivalves (Calyptogena? sp.) occur exclusively in the plume (mean abundance=0.5 individuals 1000 m^?3). Larvae from other benthic taxa known not to be endemic to Juan de Fuca vent communities, such as anthozoans, pholad clams, bryozoans and echinoderms, are less abundant in the plume than away (mean abundance=47.5 vs 16.9 individuals 1000 m^?3) at comparable depths and heights above the bottom. These results support the hypothesis that larvae of vent species are entrained into buoyant hydrothermal plumes and transported at the level of lateral spreading several hundred meters above the seafloor. The discovery of vent-associated larvae in the plume suggests that models used to predict hydrodynamic processes in the plume will also be useful for modeling larval dispersal. Advanced imaging and new molecular-based approaches will be required to resolve taxonomic uncertainties in some larval groups (e.g. certain polychaete families) in order to distinguish vent species and make comprehensive flux estimates of all vent larvae in the neutrally-buoyant plume.     

Temperature and salinity tolerances of embryos and larvae of the deep-sea mytilid mussel ��Bathymodiolus�� childressi

We examined temperature and salinity tolerances of early embryonic and larval stages of the deep-sea, cold-seep mussel ??Bathymodiolus?? childressi to determine whether they may control the dispersal depth of larvae. Salinity and temperature tolerances increased with developmental stage, but tolerance ranges were not as wide for the larvae of ??B.?? childressi as for the larvae of the related shallow-water mussel Mytilus trossulus. Normal development occurred in ??B.?? childressi from 7 to 15°C and at salinities of 35 and 45. Greater tolerance of ??B.?? childressi embryos to high than low salinities may aid development of negatively buoyant early embryos at brine seeps. Although there was a decreasing trend in survival of ??B.?? childressi larvae with increasing temperature, survival of ??B.?? childressi trochophores was not significantly different at 20°C than at the adults?? ambient temperature. Since larvae tolerate increasing temperatures as they age and seawater temperatures at 100?m depth do not exceed 20°C in months following the mussels?? spawning season, we suggest that temperature would not limit vertical migration of the veliger larvae of ??B.?? childressi into even the uppermost layer of the water column above the cold seeps in the Gulf of Mexico.     

Metabolic responses of the squat lobster (Pleuroncodes monodon) larvae to low oxygen concentration

Squat lobster populations found in the Humboldt Current System over the continental shelf from ~28 to 37°S release pelagic larvae in sub-surface cold (~11 °C) hypoxic waters. Larvae subsequently spread throughout the water column encountering both normoxic and hypoxic conditions. We analyzed some short- and long-term responses of Pleuroncodes monodon larval metabolism to hypoxia at 11 °C. Routine and postprandial aerobic respiration rates were lower in hypoxia than in normoxia for all zoeal stages. Zoea V oxyconformed, while megalopae oxyregulated down to very low oxygen concentrations. Throughout zoea I development, the rate of nitrogen (protein) accumulation in zoea I was lower, and C:N ratios were higher under hypoxic conditions than in normoxia. Citrate synthase (CS) and malate dehydrogenase (MDH) apparent specific activities (as indicators of aerobic and metabolic potentials, respectively) decreased and remained at the same level, respectively, throughout zoea I reared under hypoxic conditions. Anaerobic to aerobic potential (lactate dehydrogenase (LDH)/CS) was higher in organisms reared under hypoxia, and MDH/LDH potential ratios were characteristic of organisms tolerant to hypoxia. In spite of P. monodon zoea endurance and metabolic adaptations to decreasing oxygen tensions, intense hypoxia as such of their release site would affect their overall condition especially toward the end of the molt cycle. Our results indicate the importance of considering the interaction between environmental oxygen variability and recruitment success.     

Competition for food and other niche-related studies of three species of salt-marsh foraminifera

The niches of 3 species of salt-marsh foraminifera, Allogromia laticollaris, Rosalina leei, and Spiroloculina hyalina were assessed in the laboratory. The 3 species reproduce within the following ranges: temperature (10° to 33°C), salinity (12 to 45‰) and pH (5 to 10). Competition for food among the 3 species was evaluated. S. hyalina did not compete with other species. Intraspecific competition (crowding) appears to be an important factor limiting the reproduction of A. laticollaris. Crowding seems to have little effect on the other 2 species. The feeding of foraminifera is affected by the quality and quantity of food organisms. The feeding rate of the species tested is directly related to concentration within a range of 10² to 10⁶ cells fed. S. hyalina is a bacterial feeder. A. laticollaris is a rare species which may become locally abundant when dominant species are missing. S. hyalina is also a rare species, which can bloom where the density of bacteria is relatively high and in the absence of competing species. R. leei is a stable, conspicuous species, whose moderate numbers are relatively unaffected by physical stress and competition. Some new laboratory data on Ammonia beccarii were obtained so that this species could be compared with the other studied. Differences in niches are graphically presented.     

Shadow cinematography of fish larvae

A simple system of shadow cinematography, consisting of a small tungsten halogen lamp, 2 large biconvex lenses and a 16 mm camera, is described for recording the swimming and feeding behaviour of larval fish. The system can be used either with infra-red film to record swimming behaviour independently of ambient light intensity, or with high-resolution film to record food organisms and feeding behaviour. Small plankton organisms of 0.2 mm width can be resolved using high-resolution film. The technique has been used to record the behaviour of plaice larvae (Pleuronectes platessa L.) feeding on the nauplii of Artemia salina L. The perceptive field of the larvae extends to approximately ±60° in azimuth, ±40° in elevation and 1.5 body lengths in range.     

Dispersal of phytoplanktotrophic shipworm larvae (Bivalvia: Teredinidae) over long distances by ocean currents

Shipworms or Teredinidae may be dispersed either as adults in floating wooden objects or as pelagic larvae drifting near the sea surface. Five shipworm species, i.e., half of those having an amphi-Atlantic geographical distribution, are known also to have pelagic phytoplanktotrophic larvae which can be carried by ocean currents. From a series of 742 plankton samples taken from throughout the temperate and tropical North Atlantic Ocean, it can be shown that shipworm larvae are not uncommon in the open sea. Teredinid veligers were found in 19% of all samples taken. One species of larvae, identical in all details to that described by Rancurel (1965), is particularly prevalent and is tentatively identified as Teredora malleolus (Turton). A definitive identification will be possible only after the pelagic larvae of the other Atlantic species are known. The larvae of Teredora malleolus are found throughout the North Atlantic Gyre and the adjacent temperate and tropical seas, and from scattered records in the South Equatorial Current. Larvae of other unidentified Teredinidae species were also found. The distance that larvae may be transported depends upon the length of pelagic larval development and the velocity of the currents. From the known current velocities it can be shown that, even in a few weeks, larvae may be dispersed many hundreds of kilometers. The geographical distribution of shipworm larvae suggests that they are carried along the coasts of continents and even across ocean basins, and that this dispersal must be an important factor in the geographical distribution of the adult forms and in the maintenance of genetic continuity between populations otherwise isolated from one another.Contribution No. 2555 from the Woods Hole Oceanographic Institution, Woods Hole, Massachusetts, USA