The functional morphology and development of the alimentary tract of larval and juvenile barnacles (Cirripedia: Thoracica)

The alimentary tract of the nauplius larva of Balanus spp. consists of cuticle-lined foregut and hindgut, with intervening endodermal midgut constricted into anterior and posterior regions. The anterior midgut cells in the region of the constriction (constriction cells) secrete proteins (probably digestive enzymes). The remaining anterior midgut cells, often containing lipid droplets, form the absorptive region of the tract. Glycoprotein globules and lipid droplets within anterior midgut cells are the remants of the yolk in a pre-hatched larva, this yolk additionally supporting the larva through the non-feeding first nauplius stage. Nauplius Stages II to VI are actively feeding planktonic stages which increase in size and build up lipid reserves. These accumulated reserves support the non-feeding cyprid, first through its planktonic life and then through settlement and subsequent metamorphosis to the juvenile barnacle. Juvenile barnacles start to feed between 2 and 5 days after metamorphosis.     

Effect of delayed first feeding on larval performance of the spider crab Maja brachydactyla assessed by digestive enzyme activities and biometric parameters

The effect of food deprivation on larval performance of the spider crab Maja brachydactyla was studied in terms of survival, moulting capacity, size, weight and enzymatic activities. Five feeding treatments that differed in the initial age of first feeding larvae (fed from hatching, 2, 4 and 6 days post-hatching and unfed) were tested for 20 days. Newly hatched larvae kept without food supply lasted for 10 days and did not moult; with 50% survival observed at 6 days post-hatching. Larvae (zoea I stage) were only able to tolerate 2 days of food deprivation after the onset of exogenous feeding without their performance being compromised. Multivariate analyses suggest that digestive enzyme activities may be good indicators of the nutritional condition of larvae.     

Endogenous control of timing of metamorphosis in megalopae of the shore crab Carcinus maenas

Batches of hundreds of freshly collected megalopae of the shore crab Carcinus maenas (L.) showed persistent circatidal rhythms of moulting to the juvenile crab stage when maintained in constant laboratory conditions. Peaks of moulting occurred around expected times of high tides, with few megalopae moulting at other times. In larvae collected offshore, the highest tidally-timed peak of metamorphosis occurred during the second to fifth expected times of high tide, and metamorphosis of 50% of each batch took about 22 h or longer. In contrast, in larvae collected at the water's edge, 70% metamorphosed during the first expected episode of high tide, within 6 to 8 h after collection. However, although inshore megalopae moulted before offshore ones, the tidal timing of moulting remained unaltered whether megalopae were collected at neaps or springs, from the water's edge or farther offshore, in the presence or absence of natural substratum, and under various light–dark and salinity regimes. Metamorphosis of C. maenas megalopae around the times of high tides may enhance settlement into the upper intertidal zone. Early juveniles of the crab apparently prefer that zone as they are most abundant there and, unlike adults, do not undertake up-and-down-shore migration with tides. The present finding demonstrates, for the first time, endogenous physiological timing of circatidal periodicity in the metamorphic moult of crab megalopae, suggesting that endogenous factors, as well as exogenous ones should be taken into account in considering the process of settlement by crab megalopae. Received: 21 February 1996 / Accepted: 27 November 1996     

Effect of dissolved oxygen level on respiratory metabolism, nutritional physiology, and immune condition of southern king crab Lithodes santolla (Molina, 1782) (Decapoda, Lithodidae)

Episodes of hypoxia are common in the marine environment, and their ecological effects depend, in part, on their severity and duration. Many species of decapod crustaceans reside in areas with fluctuating oxygen regimens. Physiological mechanisms enhance the ability of these crustaceans to cope with acute episodes of hypoxia. Southern king crab, Lithodes santolla, fishery is important in the south of South America, and some data describe fishing zones with low dissolved oxygen (DO) levels (3.5 mgO₂ l⁻¹, i.e., 8.3 kPa). Our main objective was to evaluate the effect of dissolved oxygen level on respiratory metabolism, nutritional physiology, and immunological condition of L. santolla juveniles. Individual animals were exposed for 10 days to different oxygen tensions (2.1, 4.2, 8.5, 12.7, and 21.1 kPa) to quantify the oxygen consumption rate; thereafter, blood oxyhemocyanin (Hc), protein concentration, as well as hemocytes, were sampled. Freeze-dried animals were dissected, and digestive gland metabolites (glycogen, protein, glucose, cholesterol, acylglycerol, and lactate) and digestive enzyme activity (general protease, trypsin, and chymotrypsin), as well as gill lactate dehydrogenase (LDH) activity, were quantified. In the present study, Lithodes santolla showed a critical oxygen tension between 4 and 9 kPa, indicating that this crab species is more sensitive to DO than other crustacean species. Protein and Hc concentrations followed a similar pattern to that of oxygen consumption. Digestive gland glycogen and protein concentration did not change after 10 days at different oxygen exposures, but glucose, cholesterol, and acylglycerol concentrations decreased linearly and proportionally to the available oxygen in the water. As in other decapods, chymotrypsin showed over 90% of the total quantified proteases activity. Chymotrypsin activity together with total proteases and trypsin was not affected by the environmental oxygen tension. Gill LDH and digestive gland lactate followed a similar increase at lower environmental oxygen tension but dropped sharply at the lowest tension (2.1 kPa). Dissolved oxygen affected also the immune system through reduction of hemocytes. This could provide a critical window for opportunistic pathogens to become established when crabs are exposed to hypoxic conditions. L. santolla juveniles show a moderate tolerance to low oxygen availability by modifying the concentration of hemolymph proteins, mainly OxyHc, some digestive gland metabolites, and by activating the anaerobic metabolism. This allows L. santolla juveniles to inhabit temporarily low oxygen zones in the deep ocean and suggests an advantage for culture conditions.     

Vertical distributions of eggs,nauplii and copepodites of Calanus helgolandicus (Copepoda: Crustacea) in the Celtic Sea

The vertical distributions of eggs, nauplii, copepodites and adults of Calanus helgolandicus (Claus) from five oblique plankton-net hauls taken in May (1980), March and September (1981) and January (1982) at a site in the shelf sea to the south-west of the United Kingdom are described. The water depth is approximately 95 m and becomes thermally stratified during the summer months when a thermocline of ∼6 C° develops. In early spring when the water column was isothermal (∼8 °C), the development of the eggs and nauplii took place below 60 m and a single ontogenetic migration was observed between Nauplius VI (NVI) and Copepodite I (CI). As the temperature of the water increased, this migration occurred in progressively earlier naupliar stages. The eggs were distributed throughout the water column in the profile taken in early May when a 1 C° thermocline occurred between 30 to 40 m. The majority of the NI to NIV stages occurred below 40 m, with the ontogenetic migration taking place in the NIV stage; the NV and NVI stages were found above the thermocline. In September, the eggs were again distributed throughout the water column (101 490 m^-2), with a maximum number of >4 500 m^-3 occurring in the surface to 5 m depth interval. Nauplius I and II were found at all depths, demonstrating that hatching occurred throughout the water column. The ontogenetic migration in these late-summer profiles took place between the NII and NIII stages, the remainder of the nauplii being found above the thermocline in the top 20 m. This is the first time that an ontogenetic migration, similar to the developmental ascent observed in the naupliar stages of the euphausiid Euphausia superba in the deep ocean, has been shown for a copepod nauplius.     

Apparent predation on ichthyoplankton by zooplankton and fishes in nearshore waters of southern California

Plankton collected at discrete depths in Santa Monica Bay, California, USA, during January 1982 were examined for fish eggs and larvae that had been attacked or consumed by zooplankton. The bongo net remained open for only 3 min and samples were preserved within 5 min of capture. Juvenile and adult fishes that had been captured by otter trawl and preserved within 20 min of capture were examined for ingested fish eggs and larvae. Three copepods (Corycaeus anglicus, Labidocera trispinosa, and Tortanus discaudatus), one euphausid larva (Nyctiphanes simplex), one amphipod (Monoculoides sp.), and an unidentified decapod larva were found attached to fish larvae in the preserved plankton samples (attachment to 23% of the fish larvae was observed in one sample). Overall, about 5% of the white croaker (Genyonemus lineatus) larvae and 2% of the northern anchovy (Engraulis mordax) larvae had attached zooplankton predators. Most fish larvae with attached zooplankton predators were small. We found no indication of zooplankton predation on fish eggs. Few fish eggs and larvae were found in the digestive tracts of juvenile or adult fishes, and the ingested fish larvae were relatively large. The discussion considers apparent preyspecificity of the zooplankton predators as well as potential biases that may be associated with preserved samples collected by nets.     

Development of the echinothurid sea urchin Asthenosoma ijimai

Development of an echinothurid sea urchin is described for the first time; eggs of Asthenosoma ijimai (Yoshiwara) have been fertilized in the laboratory, and development has been followed light-microscopically up to the early juvenile stage. The eggs, which are orange and float, are 1.2 mm in diameter, by far the largest echinoid eggs on record. The embryos, larvae and early juveniles are lecithotrophic, and no exogenous food is needed for development. The embryonic stages include a remarkable biscuit-shaped late blastula, which has never been described for any echinoid before. The larva, with its para-arms unsupported by skeletal ossicles, is unlike the echinoid prism or pluteus and more resembles the early bipinnaria or early auricularia of asteroids and holothurians, respectively. All stages through late larvae float just beneath the surface until settlement occurs during the third and fourth weeks at 20°C. Metamorphosis from late larva to juvenile is gradual and no part of the larval body appears to be cast off.This paper is dedicated to Professor Juro Ishida on the occasion of his seventieth birthday.     

Characterisation of cysteine proteinase activities in the digestive tract of juvenile <Emphasis Type="Italic">Paragnathia formica</Emphasis> isopods,ectoparasites of estuarine fish

Juveniles of Paragnathia formica Hesse (Isopoda; Gnathiidae) are haematophagous ectoparasites, feeding on fish blood which supplies the nutrients for their development through growth and moulting, and the subsequent survival and reproduction of the free-living adults. Little is known of the mechanisms of digestion in juvenile gnathiids, and biochemical studies on the digestive tract of these interesting estuarine isopods have not been undertaken previously. Here, functionally active cathepsin-like cysteine proteinases are identified in the digestive system of juvenile praniza (fed) and zuphea (unfed) forms. The physiological pH of the digestive tract and the optimum proteolytic activities detected in praniza 3 homogenates using the cathepsin B/L, cathepsin B, and cathepsin H fluorogenic substrates, N-Carbobenzoxy-Phe-Arg-4-methoxy-2-naphthylamine (Z-phe-arg-MNA), N-Carbobenzoxy-Arg-Arg-4-methoxy-2-naphthylamine (Z-arg-arg-MNA) and Arg-4-methoxy-2-naphthylamine (H-arg-MNA), respectively, are in the acidic range (pH 5.8–6.7). Inhibition profiles against Z-phe-arg-MNA and Z-arg-arg-MNA using the cathepsin B inactivator urea, and cysteine proteinase inhibitors, support the presence of cathepsin L- and B-like enzymes. These proteolytic activities are 10–50 times higher in homogenates of praniza 3 compared with zuphea 3 forms. Histochemistry of praniza 3 sections reveals that the predominant enzyme activity towards Z-phe-arg-MNA is limited to the digestive glands during early and mid stages of digestion; later, this activity appears in the lining of the anterior hindgut. Moreover, activity towards Z-arg-arg-MNA is generally restricted to the digestive glands, and only occasionally present in the anterior hindgut. In conclusion, the digestive glands are the main site of cathepsin-like cysteine proteinase activities in P. formica juveniles; these enzymes appear to be important to the digestion of host fish blood enabling development through to the free-living adult.     

Early growth and mortality of the Caribbean king crab Mithrax spinosissimus reared in the laboratory

The berried females of the Caribbean king crab Mithrax spinosissimus (Lamarck) used in this study were collected from canals on Big Pine Key, Sugarloaf Key and Lower Matecumbe Key (south Florida, USA) on 9 August, 8 October and 15 November 1986. Viable spawns hatched as first zoeae and molted to second zoeae within ca. 10 to 12 h. Most of the larvae reached the megalopa stage 1 d later, and molted to first crab 4 to 8 d after hatching (water temperature: 27.2° to 28.8°C). Low water temperature and/or early lack of food had a negative effect not only on stage duration, but also on the size of the early crab stages. Successful molt to first crabs occurred, however, in the absence of food. The growth rate (carapace length) between molts in early crab stages varied between ca. 20 and 30%. When provided with good water exchange, stocking density could be very high (>22 500 individuals m^-2), with no increase in mortality. The highest mortality rate was recorded when the larvae molted to first crab, and the highest rates of survival were always recorded when feeding was not initiated until after 5 to 8 d after hatching. No cannibalism was observed among larvae, and cannibalism was low in early crab stages. The study indicates that to achieve viable hatches and high larval survival in rearing M. spinosissimus, a continuous and adequate supply of high-quality seawater is a prerequisite both in larviculture and in maintaining brooding females.Contribution No. 93, Department of Oceanography and Ocean Engineering, Florida Institute of Technology     

Diet and prey selectivity of scyphomedusae from Port Phillip Bay,Australia

Scyphomedusae collected from Port Phillip Bay, Victoria, Australia, between 1984 and 1986, consumed a variety of zooplankton. The percentage composition of gut contents of Cyanea capillata (Linné) in order of decreasing importance was larvaceans 31%, cladocerans 29%, fish eggs 14%, copepods 11%, hydromedusae 9%, and ascidian tadpoles 3%. The percentage composition of gut contents of Pseudorhiza haeckeli Haacke was fish eggs 41%, copepods 33%, larvaceans 8%, cladocerans 4%, crab zoea 4%, and decapod larvae 1%. Both species of scyphomedusae showed strong positive selection for fish eggs and yolk-sac larvae, and negative selection for other prey items. When fish eggs were omitted from the selectivity analyses, C. capillata showed positive selection for amphipods, decapods, crab zoea, Podon spp., larvaceans and ascidian tadpoles, and negative selection for Evadne spp. and all copepod taxa. Pseudorhiza haeckeli showed positive selection for amphipods, decapod larvae, crab zoea and cladocerans, and negative selection for cirripede larvae, larvaceans and hydromedusae. Amongst copepods, P. haeckeli showed positive selection for calanoid and harpacticoid copepods and negative selection for cyclopoid copepods.     

Patterns of larval and early juvenile growth in a semiterrestrial crab, Armases angustipes (Decapoda: Sesarmidae): comparison with a congener with abbreviated development

In a semiterrestrial and estuarine tropical crab, Armases angustipes Dana (Grapsoidea: Sesarmidae), changes in biomass (measured as dry mass, W; carbon, C; nitrogen, N; and hydrogen, H; per individual) and relative elemental composition (C, N, H, in percent of W; C:N mass ratio) were studied during development from an early egg stage through hatching, the complete larval phase, metamorphosis and the first juvenile crab stage (CI). In the megalopa and CI, growth was measured also within the moulting cycle, and biomass and elemental composition were determined in cast exuviae. From an early egg stage to the freshly hatched larva, A. angustipes lost about 20% of W, 29% of C, 5% of N and 32% of H. Proportionally higher losses in C than in N were reflected also in a significantly decreasing C:N mass ratio (from 5.02 to 3.74). These results indicate that lipids mobilised from yolk reserves represented the principal metabolic substrate for embryonic energy production, while proteins were catabolised at a much lower rate. The present data of growth and exuviation are compared with previously published data from a congener, A. miersii Rathbun, which has an abbreviated and facultatively lecithotrophic mode of larval development (with three instead of four zoeal stages; stages I and II in principle independent of food). When growth is measured as an increase in the final (premoult) biomass of successive developmental stages, both species show an exponential pattern. Within the moulting cycles of the megalopa and the first juvenile, both species show parabola-shaped growth curves, with a rapid biomass increase in postmoult and intermoult stages, and losses in the premoult phase. Thus, the two Armases species show, in general, similar patterns of larval and early juvenile growth. However, the initial size of eggs and larvae is about four times larger in A. miersii, and its biomass remains higher throughout the period of larval and early juvenile development. A. angustipes is able to partially make up for this difference, as it has an additional zoeal stage, and its megalopa and CI stages show higher relative biomass increments (in percent of initial values). Due to this compensatory growth pattern, A. angustipes reaches in its CI stage about half the biomass of a juvenile A. miersii. When exuvial losses of megalopae and juveniles are compared between these two species, A. miersii shows higher biomass losses per individual (corresponding with its larger size), but lower relative losses (C, N, H, in percent of late premoult body mass or in percent of previously achieved growth increments). Differences in larval and early juvenile growth and in the exuvial losses of megalopae and juveniles of these two congeners are discussed in relation to their differential ecology, life history and reproductive strategy.     

Effects of the juvenile hormone mimic ZR-512 (Altozar®) on larval development of the mud-crab Rhithropanopeus harrisii at various cyclic temperatures

Effects of the juvenile hormone (JH) mimic hydroprene (Altozar^®: ZR-512), which exhibits high activity against Lepidoptera, were studied on the larval development of the mud-crab Rhithropanopeus harrisii (Gould) (Brachyura: Xanthidae). Larvae reared in 20 S at 3 cycles of temperature of 20° to 25°C, 25° to 30°C and 30° to 35°C, were exposed to 0.01, 0.1 and 0.5 ppm hydroprene from hatching to the first crab stage. Larvae were also exposed to 0.1 and 0.5 ppm hydroprene only from the megalopa stage to the first crab stage. When larvae were treated with hydroprene throughout larval life, survival was significantly reduced with increasing concentrations of the compound at all temperature cycles. Synergistic effect between hydroprene and temperature on survival of zoeal larvae was not observed. On the average there was 11% less survival in the zoeal stages at the 0.01 ppm concentration. of hydroprene than in the control, an additional reduction of 13% occurred at 0.1 ppm, and finally there was a further decrease of 46% at 0.5 ppm hydroprene. Significant decrease in survival in the megalopa stage occurred only in the 0.5 ppm concentration of hydroprene at the lowest temperature cycle when larvae were exposed to the compound from hatching. When larvae were treated with hydroprene only within the megalopa stage, a significant reduction in survival was not observed. First-stage zoeae were the most sensitive of the larval stages to hydroprene. Duration of zoeal development was significantly delayed at 0.5 ppm hydroprene at the two lower temperature cycles, whereas in the megalopa stage the delay began at the 0.1 ppm level at all 3 temperature cycles when larvae were exposed to hydroprene from hatching. A significant delay was also observed at 0.1 ppm hydroprene at the two lower cycles when larvae were exposed to hydroprene only in the megalopa stage; at 30° to 35°C a significant delay was observed only at the 0.5 ppm level. The results show that metamorphosis to the first crab stage was not inhibited at the 0.5 ppm level of hydroprene or lower. Reduction in survival and increase in duration of larval development were presumably related to stress conditions caused by hydroprene. The results also suggest an interaction between temperature and hydroprene on survival of megalopa larvae and duration of larval development.     

Larval development of <Emphasis Type="Italic">Lightiella magdalenina</Emphasis> (Crustacea,Cephalocarida)

The larval development of the newly discovered cephalocarid from Mediterranean sea, Lightiella magdalenina, was analysed using light and scanning electron microscopy. Twenty-nine larval specimens, divided into 15 metanaupliar and 2 juvenile stages of development, were found. The first six metanaupliar stages had an even number (6, 8, 10, 12, 14 and 16) of trunk segments including telson. The condition of 20 trunk segments, typical of the adult, was reached after another 4 stages by the addition of a single segment per stage. At this tenth stage, the larvae had an incomplete number of trunk limbs. Another five stages were needed to complete the cephalic appendage development, passing from the metanaupliar to the juvenile stage, characterized by loss of the naupliar enditic process of the second antenna. Trunk limbs development was completed during the last two juvenile stages. According to the ontogenetic data reported for Hutchinsoniella macracantha and Lightiella incisa, these seventeen stages probably do not represent the complete developmental series. Nevertheless, they allow us to clarify the main features of cephalocarid ontogeny and show specific differences in the development of both the trunk segments and appendages.     

The predatory mite Phytoseiulus persimilis manipulates imprinting among offspring through egg placement

A crucial question with respect to imprinting is how animals ensure that kin imprint on kin but not on non-kin. Imprinting takes place in a sensitive ontogenetic phase, usually in an early period of life or when offspring are produced, at which time the recipient imprints on the first referents met. In the predatory mite Phytoseiulus persimilis, imprinting among immature individuals happens in the larval stage immediately after hatching. I tested the hypothesis that adult P. persimilis females manipulate offspring imprinting by influencing the likelihood of encounters among recipients and referents via egg placement and egg aggregation. I conducted two experiments, one of which addressed imprinting and cannibalism, and the other addressed egg placement and egg aggregation. The imprinting experiment suggests that larvae imprint on any conspecific individual met in a sensitive ontogenetic phase and later on treat this individual as kin, irrespective of relatedness. After molting to protonymphs, imprinted individuals preferentially cannibalized unfamiliar to familiar larvae. Irrespective of familiarity, kin were cannibalized earlier than non-kin, suggesting the involvement of self-referent phenotype matching. The egg-placement experiment provides evidence that females adjust the aggregation level of their own eggs according to the degree of relatedness to present eggs from a previously ovipositing female. Both experiments in concert suggest that egg placement is a maternal strategy influencing imprinting among immature individuals. Apart from avoiding kin cannibalism, egg placement and imprinting by larvae may have relevance to other behaviors influenced by kin recognition, such as mate choice, prey-patch choice and dispersal.Communicated by L. Simmons     

Effects on larval crabs of exposure to algal toxin via ingestion of heterotrophic prey

We tested whether ingesting toxic algae by heterotrophic prey affected their nutritional value to crab larval predators, using toxic algal strains that are either ingested directly by larval crabs or rejected by them. Ingestion of toxic strains of the dinoflagellates Alexandrium andersoni and A. fundyense by the rotifer Brachionus plicatilis was confirmed. Rotifers having ingested either algal type for five days were fed to freshly hatched larvae of three crab species, with larval survival and stage durations determined. For both algal/rotifer treatments in all three crab species, larvae fed algae directly died during the first zoeal stage, while those fed rotifers that had been fed either algal strain survived to the experiment’s end (zoeal stage 3). Survival was lower, and stage duration longer, for larvae fed rotifers cultured on toxic algae when compared to those fed non-toxic algae. The role of toxic algae in the planktonic food web may be influenced by its direct or indirect ingestion by larval crabs.     

Physiological and biochemical changes during lecithotrophic larval development and early juvenile growth in the northern stone crab,Lithodes maja (Decapoda: Anomura)

Larvae of the northern stone crab, Lithodes maja L., were reared in the laboratory from hatching to the second crab stage. complete larval development (at constant 9°C) lasted about 7 wk, invariably consisting of three pelagic zoeal stages and a semibenthic Megalopa; only two zoeal stages have been described in the literature. All larval stages are lecithotrophic. First feeding was consistently observed only after metamorphosis, in the first juvenile crab stage. In short intervals (every 1 to 5 d), developmental changes in biomass, B (expressed as: dry weight, W; carbon, C; nitrogen, N; hydrogen, H) and oxygen consumption (respiration, R) were measured in larvae and early juveniles; additionally, protein and carbohydrates were measured, but only in the zoeal stages and early Megalopa. Unusually high C contents (varying between 56 and 61% of W in eggs and freshly hatched Zoea I larvae from 12 different females) and high C:N weight ratios (8 to 11) indicate enhanced initial lipid stores, which are utilized as the major metabolic substrate during both embryonic and lecithotrophic larval development. Predominant degradation of lipids is shown indirectly; the C:N ratio decreased significantly, from 10 (at hatching) to 6 (at metamorphosis), while larval protein decreased only little, from ca. 55% of W (at hatching) to 48% (in the Megalopa). From hatching to metamorphosis, about 27% of the initially present W, 48% of C, 18% of N, and 52% of H were lost. This decrease in larval biomass can be described as an exponential function of development time. The major part of these losses were associated with metabolic energy requirements, while exuvial losses were comparably small. In each of the zoeal stages, only about 1 to 2% of late premoult (LPM) B was shed with the exuvia. The Megalopa, which produces a much thicker, calcified exoskeleton, lost 20% of LPM W, but only 5 to 8% of organic constituents (C, N, H). Much higher exuvial losses were measured in the Crab I stage (51% in W, 21% in C, 5% in N, and 7% in H). Maximum respiration was found in the actively swimming zoeal stages, a minimum in the predominantly benthic, mostly inactive Megalopa. The Crab I stage exhibits also a sluggish behaviour and low R, in spite of beginning food uptake and growth. Immediately after metamorphosis, the juvenile crab gained rapidly in W, in particular in its C fraction. A transitorily steep increase in the C:N ratio indicates a replenishment of partially depleted lipid stores, but also a rapid initial increase of inorganic C in the heavily calcified exoskeleton. Instantaneous rates of growth, assimilation, and net growth efficiency (K ₂) were high during the initial (postmoult) phase in the first juvenile crab stage (C-specific growth rate: 6% d^-1; K ₂:70%), but decreased towards zero values during laterstages of the moulting cycle; metabolism remained practically constant during the Crab I stage. Entirely lecithotrophic larval development from hatching to metamorphosis in L. maja is considered an adaptation to seasonally short and limited planktonic food production in subarctic regions of the northern Atlantic.     

Long-term effects of water-soluble fractions of Kuwait crude oil on the larval and juvenile development of the mud crab Eurypanopeus depressus

Larvae of the mud crab Eurypanopeus depressus (Smith) were reared in various concentrations of the water-soluble fraction of Kuwait crude oil. The 48-h TL_m (median tolerance limit) for Zoea Stage I was approximately 10 ppm total dissolved hydrocarbons and that for Zoea Stage II approximately 17 ppm. Chronic toxicity of more dilute solutions (4.3 and 8.7 ppm) was assessed independently for each larval stage and for subsequent developmental stages through Crab Stage 5. In the group continuously exposed to oil from hatching, there was differential mortality relative to controls in every larval stage and increased duration of intermolt periods was observed at every stage through Crab Stage 5. Mortality in groups not exposed until larvae had reached Zoea Stages III or IV was not greater than controls, suggesting that toxicity to advanced larval stages may be related to accumulation of toxic compounds by the larvae. Neither concentration (4.3 or 8.7 ppm) of crude oil caused increased mortality among juvenile crab stages regardless of the time of initial exposure. Increased occurrence of an extra and morphologically abnormal megalopa stage was associated with exposure to the crude oil. This has not been reported before.Communicated by I. Morris, West Boothbay Harbor     

Larval settlement and metamorphosis in the sand crab Emerita talpoida (Crustacea: Decapoda: Anomura)

The effects of various potential cues on the survival and time to metamorphosis of larvae of the sand crab Emerita talpoida (Say) were examined. Zoeal duration, which ranged from 25 to 43 d, was not correlated with subsequent megalopal survival but had a weak, negative correlation with the duration of the megalopal stage. Neither food, sand nor water from containers with adults directly affected megalopal survival or duration. In a second experiment, the presence of sand both increased the probability of survival and shortened the duration of late zoeal stages. This is the first experimental evidence for delayed zoeal metamorphosis in a decapod. Settlement of E. talpoida appears to occur primarily during the zoeal phase, not at the megalopal stage.     

Maternal provisioning for larvae and larval provisioning for juveniles in the toxopneustid sea urchin <Emphasis Type="Italic">Tripneustes gratilla</Emphasis>

Lipid and protein biochemistry of eggs (84 μm in diameter), embryos and early larvae of the tropical echinoid Tripneustes gratilla (Linnaeus 1758) were quantified to determine how maternal provisions are used to fuel development of the echinopluteus. The eggs contained a mean of 30.82 ng lipid and 87.32 ng protein. Energetic lipids were the major lipid component (55.52% of total lipid) with the major class being triglyceride (TG: mean 15.9 ng, 51.58% of total). Structural lipid was dominated by phospholipid (PL: mean 11.18 ng, 36.26% of total). Early embryogenesis was not a major drain on egg energetic lipid and protein. Development of the functional feeding larva used ca. 50% of initial egg energetic lipid and most of this was TG. Maternal TG was still present in the 8-day echinoplutei and it was estimated that this energetic lipid would be depleted in unfed larvae by day 10. There was no change in PL. In a separate experiment lipid biochemistry of rudiment stage larvae and early developing juveniles were quantified to determine how lipids are used during metamorphosis. Fed larvae accumulated lipid (mean 275.49 ng) with TG and PL being the major energetic and structural lipids, respectively. Larval lipid stores were not appreciably depleted by metamorphosis and so were available for the early benthic stage juvenile. Juveniles started their benthic existence with 314 ng total lipid (TG: mean 46.84 ng, 14.9% of total, PL: mean 137.51 ng, 43.67% of total). Nile Red histochemistry and histology showed that the stomach serves as a nutrient storage organ and, that lipid stores accrued by larvae sustain developing juveniles for up to 4 days post settlement. Triglyceride supported both non-feeding stages of development and the prefeeding larval and perimetamorphic benthic stage. In this first study of lipid stores in settlement stage echinoderm larvae, we show that T. gratilla larvae sequester the same major energetic lipid (TG) to support the early juvenile that the female parent provided them to fuel early development.