Diel trophic interactions between vertically-migrating zooplankton and their fish predators in an eelgrass community

Diel changes in the composition of crustacean zooplankton and the diets of fish predators from an intertidal eelgrass flat were monitored concurrently. The zooplankton is characterized by two major components. The obligate zooplankters (holoplanktonic calanoid copepods and meroplanktonic decapod larvae) appear to exhibit vertical migration, being present in higher densities near the surface of the water column at night. The facultative zooplankton (amphipods and ostracods) are benthic during the day, but move up into the water column at night. Planktivorous midwaterdwelling fish consume calanoid copepods and decapod larvae during the day and cease feeding or switch their diet to amphipods at night. Benthic-dwelling fish consume some amphipods during both day and night. The factors important in prey selection by fish and the functional significance of vertical migration in both components of the zooplankton are discussed in the light of the changing patterns of fish predation.This paper is Publication No. 183 in the Ministry for Conservation of Victoria, Environmental Studies Series.     

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.     

Predation on zooplankton by the benthic anthozoans Alcyonium siderium (Alcyonacea) and Metridium senile (Actiniaria) in the New England subtidal

The Alcyonacean octocoral Alcyonium siderium Verrill and the sea anemone Metridium senile (L.), the only common perennial zooplanktivores on shallow (16-m depth) subtidal rock walls in much of northern New England, USA, are of similar heights and overlap in their habitat and microhabit distributions. The coelenteron contents of both species were sampled at four-hour intervals over a diel cycle and were compared to zooplankton available in the water at 1 to 5 cm from the rock wall, the height at which the cnidarians held their feeding tentacles. Prey in coelenterons of A. siderium were significantly smaller (means of 256 to 345 m), and those in coelenterons of M. senile were equal to or slightly greater in length (means of 415 to 1006 m) than the available zooplankton. The diets of A. siderium and M. senile differed significantly from each other and from the available zooplankton. A. siderium showed strong positive electivites for ascidian larvae and for foraminiferans, and strongly negative electivities for most crustaceans. M. senile had strong positive electivities for barnacle cyprids, ascidian larvae, and gammarid amphipods, and strong negative electivities for invertebrate eggs, foraminiferans, calanoid and harpacticoid copepods, and ostracods. Electivities may reflect tentacle avoidance or escape by motile prey as well as predator preference. Substratum-associated organisms (e.g. demersal crustaceans, larvae of benthic invertebrates) were the most common items in the diets of both species, suggesting a tight benthic food web, similar to the situation for coral reef anthrozoans which rely on reef-generated zooplankton. A. siderium ate large numbers of ascidian larvae which, as benthic adults, compete for space with A. siderium and can overgrow small colonies. Predation on the larvae of a competing species may alleviate competition by decreasing the competitor's recruitment.     

Chemical defense and aposematic coloration in larvae of the ascidian Ecteinascidia turbinata

In the coastal waters of Florida (USA) tadpole larvae of the colonial ascidian Ecteinascidia turbinata contain chemicals which make them unpalatable to planktivorous juvenile pinfish Lagodon rhomboides. Experiments demonstrate that the bright organe color of E. turbinata tadpoles is aposematic. Fish that have recently tasted larvae of E. turbinata will not attack the palatable tadpoles of Clavelina oblonga when the latter are dyed organe to resemble larvae of E. turbinata. Tadpoles of E. turbinata that have been mouthed and rejected by fish generally survive to complete a normal metamorphosis. Individual selection explains the evolution of aposematic coloration in E. turbinata better than kin selection. The identity of the defensive chemical is unknown. The unpalatable substance in larvae of E. turbinata is removed by dialysis, indicating that it has a molecular weight less than 14000 d. Larvae are not acidic, nor is the active substance denatured by doiling.     

Seasonal development of scyphozoan medusae and the predatory impact of Aurelia aurita on the zooplankton community in the Bornholm Basin (central Baltic Sea)

The annual cycle of abundance and distribution of the scyphozoan medusae Aurelia aurita and Cyanea capillata was studied in the Bornholm Basin (central Baltic Sea) in 2002. Seasonal changes in prey composition and predatory impact were investigated by analyzing stomach contents. A. aurita occurred from July to November, with a maximum mean abundance of 2.3 ind. per 100 m³ in August, whereas C. capillata was caught in much smaller numbers from July to September. No ephyrae of either species were found; therefore, advection of medusae from the western Baltic Sea is assumed. From July to October, ~80% of A. aurita medusae was distributed in the upper 20 m above the thermocline, whereas C. capillata occurred only in the halocline below 45 m. A. aurita did not migrate vertically and fed mainly on the most abundant cladoceran species Bosmina coregoni maritima. Further prey organisms were the cladocerans Evadne nordmanni and Podon spp., mollusk larvae and copepods. Copepod nauplii and copepodite stages I–III were not eaten by the medusae, neither were fish eggs and larvae used as prey. Based on mean medusa and zooplankton abundance from the upper 20 m, the predatory impact was very low. In August, when mean abundance of A. aurita was highest, only 0.1% of the copepod and 0.5% of the cladoceran standing stock were eaten per day. However, in regions with higher medusa or lower zooplankton abundance, up to 7.9% of the cladoceran standing stock was consumed per day. Hence, A. aurita did not regulate the zooplankton community in the Bornholm Basin, and fish larvae did not suffer from competition with and predation by the medusae.Communicated by O. Kinne, Oldendorf/Luhe     

Digestive enzymes in the ontogenetic stages of the southern king crab, Lithodes santolla

The early ontogenetic stages of the sub-Antarctic king crab Lithodes santolla were analyzed for the presence and activities of a set of important digestive enzymes. The eggs and non-feeding larvae (zoea I-III, megalopa) showed high activities of esterases, phosphatases, and exopeptidases indicating the enzymatic ability to utilize endogeneous yolk reserves. SDS-PAGE showed a continuous decrease of proteins or proteids in the range of 59–81 kDa during ontogenetic development from the eggs through the zoeal stages to the first juvenile crab stage, CI. This reduction reflects the degradation of storage compounds during lecithotrophic larval development. Activities of the endopeptidases, trypsin and chymotrypsin, were low in eggs and larvae but increased significantly in the first juvenile crab stage. These enzymes typically facilitate the first steps of proteolysis in the extra-cellular spaces of the midgut gland and in the stomach. Their scarcity indicates that the larvae of L. santolla are physiologically not prepared to digest external food. This ability seems to appear first in the CI stage. Extracts of juvenile midgut glands and the gastric fluids of adults showed high activities of a variety of digestive enzymes including phosphatases, carbohydrases, as well as endo- and exopeptidases. High activities of digestive enzymes in adults may compensate for scarce food supply and rate-limiting low temperatures in the predominantly sub-Antarctic habitats of L. santolla.     

Abundance,distribution and prey composition of scyphomedusae in the southern North Sea

The annual cycle of abundance and distribution of the scyphozoan medusae Aurelia aurita, Cyanea lamarckii, C. capillata and Chrysaora hysoscella were studied in the southern North Sea in 2004 and 2005. Three different patterns of seasonal occurrence of medusae were distinguished: (1) the early occurring C. lamarckii (February–August), (2) C. capillata and A. aurita (April–August) and (3) the late appearing C. hysoscella (July/August–September). Cyanea lamarckii was the most frequently encountered species in this study; its highest mean abundance was 1.8 ± 2.7 ind. 100 m⁻³. The prey spectra of C. lamarckii, C. capillata and C. hysoscella contained several copepod and other crustacean species and thus make them potential competitors with fish larvae. Medusae in this study also consumed fish eggs and larvae, including clupeids, in all months analysed. Although peak spawning of sprat (Sprattus sprattus) coincides with the maximum abundance of medusae (May–June) the relative low abundance of all medusae species in this study makes jellyfish predation unlikely to be a factor controlling sprat recruitment in the time frame investigated.     

Feeding rates and prey selectivity of planktonic decapod larvae in the Western English Channel

Meroplankton are seasonally important contributors to the zooplankton, particularly at inshore sites, yet their feeding ecology is poorly known relative to holoplankton. While several studies have measured feeding in decapod larvae, few studies have examined the feeding rates of decapod larvae on natural prey assemblages throughout the reproductive season. We conducted 8 feeding experiments with Necora puber, Liocarcinus spp. and Upogebia spp. zoea larvae collected from the L4 monitoring site off Plymouth (50°15.00′N, 4°13.02′W) during spring–summer 2009 and 2010. This period spanned moderate-to-high food availability (0.5–1.6 µg chl-a L^?1), but a great range in food composition with small cells <20 µm dominating in 2010. Daily rations averaged 17, 60 and 22 % of body C for the 3 respective decapod species. Clearance rates differed according to prey type, and all 3 decapod genera showed evidence of selection of dinoflagellates. Importantly, small cells including nano- and pico-plankton were ingested, this being demonstrated independently by flow cytometric analysis of the feeding experiments and molecular analysis. PCR-based analysis of the haptophyte portion of the diet revealed ingestion of Isochrysis galbana by decapod larvae in the bottle incubations and Isochrysis galbana and Phaeocystis globosa by decapod larvae collected directly from the field. This study has shown that pico- and nano-sized plankton form an important supplement to the diverse and variable diet of decapod larvae.     

Food and feeding dynamics of the larval Antarctic fishNototheniops larseni

The feeding dynamics of larvae of the Antarctic fishNototheniops larseni were analyzed from data collected over three years in Bransfield Strait and adjacent waters (Antarctica). Seasonal feeding was examined from 1977/1978 (November–March). The diel feeding cycle was investigated during a 96 h station established in February 1976, while food selection was analyzed using larvae and zooplankton samples collected in February 1982. Hatching occurs in early spring, and larvae fed on eggs of calanoid copepods and on cyclopoid copepods. Copepod eggs were the principal food near the pack ice, and cyclopoids in open waters. Cyclopoids were the staple food in summer. Eggs of the Antarctic krillEuphausia superba were ingested selectively and formed major portions of the larval summer diet in neritic (Joinville Island) and oceanic (Elephant Island) spawning areas ofE. superba. In the fall, copepods predominated in the diets. Most abundant and most frequently ingested prey in summer and fall wereOncaea spp. Feeding commenced at dawn and continued at least until dusk. Krill eggs were taken chiefly during morning hours and egg incidence declined during the day, suggesting that eggs were ingested soon after spawning. Prey size at the onset of feeding was estimated as 0.130 to 0.330 mm. Size-selective feeding was evident in small larvae, while in larger larvae median prey length remained constant. High feeding incidence among yolk-sac larvae in spring, high overall feeding incidence in summer, and size-selective foraging of small larvae suggested favorable feeding conditions in the 1977/1978 season. Yolk-absorption times in Antarctic fish larvae vary on a scale of weeks and may be further retarded due to early feeding. Hence, year-to-year variability of yolk incidence inN. larseni indicated variable biotic environments of early feeding larvae rather than temporal shifts of hatching periods. As hatching periods are constant between years in contrast to the variable retreat of the pack ice and subsequent onset of the production cycle in space and time, maternal yolk reserves are probably utilized to compensate for such variations.     

Plankton community dynamics of the central Great Barrier Reef Lagoon: Analysis of data from Ikeda et al.

Plankton data collected by Ikeda et al. (1980) from the central region of the Great Barrier Reef, and spanning two years (1976 through 1978) of zooplankton records, have been analyzed extensively for spatial and temporal patterns. Estimates of net zooplankton (including chaetognaths, copepods, and larvaceans) and microzooplankton (juvenile copepods, encompassing nauplii and copepodites, and ciliates) were assessed at three stations across the 60 km lagoon. Temperature, salinity, and chlorophyll a were also measured. A cross-lagoonal gradient was identified in the plankton, concurring with results of related surveys of benthic taxa, such as scleractinian corals, soft corals, macro-algae, fish, sponges, crinoids, etc. Two associations of net zooplankton were identified. The first was associated primarily with the inner lagoon; the second with the outer lagoon. The inshore association was characterized by higher abundances of almost all net zooplankton taxa, particularly chaetognaths, copepods, polychaetes, decapods, and meroplanktonic larvae as well as higher concentrations of chlorophyll a. This inshore association wove back and forth across the lagoon through time, dominating the lagoon entirely during periods of high river discharge, reaching the mid-shelf platform reefs in this region, and sometimes being entirely absent during dry periods. Both seasonal and annual peaks in plankton abundance were generally linked with degree of runoff. Summer/autumn peaks of abundance were evident in chaetognaths, copepods, and larvaceans while annual variation was detected in the former two as well as in chlorophyll a concentrations. Depth stratification was noted in juvenile copepods and chlorophyll a concentrations at the center of the lagoon, with higher abundances recorded in deeper waters. The central Great Barrier Reef lagoon was found to be typical of other tropical coastal waters where plankton community dynamics are controlled primarily by physical factors. We suggest that any substantial changes in river discharge in this area will affect plankton production.A.I.M.S. Contribution No. 242     

Studies on portunid crabs from the Eastern Pacific. II. Significance of the unusual distribution of Euphylax dovii

Euphylax dovii Stimpson (Brachyura: Portunidae: Podophthalminae), a tropical Eastern Pacific swimming crab, has distinctive morphological adaptations for pelagic existence. Crabs in collections from the open ocean had a sex ratio approximating 1:1, with no crabs bearing eggs. Samples from the continental shelf of Colombia contained thousands of females, mostly ovigerous, but no males. Egg attachment has posed a major problem in the evolution of decapod crustaceans, and the two genera of portunid crabs thus far observed cannot attach eggs unless females can bury partly in soft sediments. This suggests that mated E. dovii females must migrate into shallow shelf waters to encounter sediments necessary for spawning. The high energetic cost of swimming while carrying eggs and the presence of abundant food for larvae are factors favoring residence of females in shelf waters until hatching is complete.     

A commensal relationship between the scyphozoan medusae<Emphasis Type="Italic"> Catostylus mosaicus</Emphasis> and the copepod<Emphasis Type="Italic"> Paramacrochiron maximum</Emphasis>

The copepod Paramacrochiron maximum was found in high numbers (up to 5,675 copepods/medusa) on the oral arms of the scyphozoan Catostylus mosaicus. This association was considered to be commensalism for the following reasons: P. maximum (Lichomolgidae) was abundant on the medusae (approximately 805 copepods/kg of medusae) and very rare in the water column (approximately 5.99×10^-4 copepods/kg of water); copepodites and adults of the symbiont were present on the host; the copepods were on the medusae both day and night, at different times (nine occasions between March 1999 and May 2000) and different locations (Botany Bay and Lake Illawarra, NSW, Australia). Over 40 taxa of plankton were found on the oral arms of C. mosaicus (including protists, cnidarians, polychaetes, molluscs, a wide range of holoplanktonic and meroplanktonic crustaceans, chaetognaths and fish eggs). These taxa were abundant in the water column and we concluded that they were prey. Symbiotic amphipods and carangid fishes were found with medusae. We conclude that there is a symbiotic association between P. maximum and C. mosaicus and care should be taken not to confound these copepods with the prey of C. mosaicus. Poecilostomid copepods are well known for consuming mucus and feeding is likely to be a major reason for the association.Communicated by G.F. Humphrey, Sydney     

Larval growth in the estuarine crab Chasmagnathus granulata: the importance of salinity experienced during embryonic development, and the initial larval biomass

The importance of salinity experienced during embryonic development and initial larval biomass on larval growth was studied in the South American estuarine crab Chasmagnathus granulata. Ovigerous females were maintained at three salinities (15, 20, and 32‰) from egg laying to hatching of zoea l. Larvae from all treatments were reared under constant conditions of photoperiod (12∶12), temperature (18°C), and salinity (first instar at 20‰, subsequent instars at 32‰). Biomass was measured as dry weight, carbon, and nitrogen content per individual at egg laying, hatching of zoea l, premoult zoea l, and zoea 4, and in 8-day-old megalopa. From hatching to premoult zoea 4, biomass was higher for larvae from prehatching salinities of 15 and 32‰. There was a significant positive correlation between biomass at hatching and at premoult zoea l and zoea 4. Accumulated biomass during zoeal stages tended to be higher for larvae from broods with higher biomass at hatching, although this trend was not always significant. Zoea 4 either directly metamorphosed to megalopa or moulted to zoea 5, following, respectively, a short or long developmental pathway. The proportion of zoea 4 that followed the long pathway was negatively correlated with biomass of zoeal stages. Biomass at hatching was correlated with biomass of megalopae developed through the short pathway, although it was not correlated with the accumulated biomass at this stage. Megalopae developed through the long pathway (i.e. metamorphosed from zoeae 5) had higher biomass than those from the short pathway. The present results suggest that prehatching salinity and initial egg and larval biomass can be very important for larval growth. Published online: 9 August 2002     

Providing a common diet to different marine decapods does not standardize the fatty acid profiles of their larvae: a warning sign for experimentation using invertebrate larvae produced in captivity

Larval decapods are commonly produced in captivity and employed in experiments to evaluate interspecific physiological and biochemical differences. Currently, it is still unknown if different decapod species provided a common diet and exposed to identical abiotic conditions produce newly hatched larvae (NHL) with similar fatty acid (FA) profiles. This study analyzed the FA composition of NHL from five marine shrimp species (Lysmata amboinensis, L. boggessi, L. debelius, L. seticaudata and Rhynchocinetes durbanensis) fed a common diet and stocked at constant temperature. FA profiles of NHL differed significantly within and among genera. NHL from species unable to molt from zoea I to zoea II in the absence of food (L. amboinensis, L. debelius and R. durbanensis) displayed the lowest FA contents. Researchers must be aware that providing a common diet to different species, even if closely related, may not standardize the FA profile of NHL produced in captivity.     

Mesozooplankton resting eggs in the Baltic Sea: identification and vertical distribution in laminated and mixed sediments

Benthic resting eggs of rotifers, cladocerans and calanoid copepods were studied at two sites with different hydrographical and sediment properties off the SW coast of Finland, the northern Baltic Sea between 1991 and 1993. Vertical distribution of the resting eggs in the sediment was studied down to 10 cm depth, and hatching of the eggs extracted from different 1-cm thick sediment layers was experimentally tested. The larvae of eight mesozooplankton taxa emerged from the eggs: Calanoida (mostlyAcartia bifilosa), cladoceransBosmina spp.,Daphnia spp.,Evadne nordmanni, andPodon spp. (mostlyP. polyphemoides), and rotifersSynchaeta spp.,Keratella spp. andAsplanchna priodonta. Some calanoid and cladoceran eggs hatched after 7 mo storage of the sediment at 3°C. A few calanoid and cladoceran eggs hatched from the 8 to 10 cm deep sediment layer. Their age, estimated by means of¹³⁷Cs analysis, was ca. 8 to 10 yr. Vertical distribution of the benthic eggs reflected the different sediment structures of the study sites. The eggs of all zooplankton taxa were more stratified in the laminated sediment than in the disturbed sediment. We suggest that the intensity of deep water mixing is the key factor for regulating the vertical distribution and hatching probability of mesozooplankton resting eggs, since it largely determines the stratification of the sedimenting material, distribution of oxygen within the sediment, and the living conditions of benthic animals causing bioturbation.     

Neue ergebnisse über die aufzucht von Carcinus maenas im laboratorium

The shore crab Carcinus maenas was reared in the laboratory from egg deposition to sexual maturity. Special enclosures were developed for cultivation of the larvae. Food and temperature proved to be the most important exogenous factors for rearing success. Fresh Artemia salina nauplii were the only food suitable for all larval stages. The following rearing temperatures proved most successful during larval development: (1) embryonic development, 10°C; (2) zoea stages, 15°C; (3) megalopa stage, 17.5°C. The larvae hatch preferably in darkness when reared under short-day conditions.     

Temporal variability in functional responses and prey selectivity of the pelagic mysid, Mysis mixta, in natural prey assemblages

Predation rates and prey selection of the pelagic mysid shrimp, Mysis mixta, were studied experimentally in the northern Baltic Sea in 1998 during their most intensive growth period, from June to October. Functional responses during 5 months were determined by providing the mysids with a natural zooplankton assemblage, diluted to several different concentrations. The results show that ingestion rate increased, along with mysid growth, from early summer to autumn and that saturation level was reached between 400 and 500 μg C l⁻¹. Ingestion rates increased with increasing prey concentration, and sigmoidal curves explained mostly the variation in ingestion rates (explanatory levels of 86–97%). Prey selection was evident in June, July and August, though weaker during the latter 2 months. Selection differed between the studied months but, generally, copepods were more positively selected than cladocerans. Rotifers were the main prey during June and July, when mysids were small, while larger mysids fed on copepods and cladocerans. Of the copepods, Eurytemora affinis was a truly selected species. This study shows that mysids feed on many zooplankton taxa and that they undergo ontogenetic diet shifts. Received: 19 July 2000 / Accepted: 19 October 2000     

Energetics,moult cycle and ecdysteroid titers in spider crab (Hyas araneus) larvae starved after the D0 threshold

Zoea I larvae of Hyas araneus L. (Decapoda: Brachyura: Majidae) were dredged in January 1986 from the German Bight and reared in the laboratory at constant 12°C, until they reached the transition of stages C/D₀ of the moult cycle (4 d after hatching). This developmental stage had previously been found to correspond with the point of reserve saturation (PRS) which allows autonomous (food-independent) development through the rest of the moult cycle and hence, was termed the D₀ threshold. One part of the larvae was continually fed (control), another group was starved from the D₀ threshold until moulting to the zoea II instar. In these two experimental groups, as well as in the two groups of zoea II larvae obtained from the different feeding conditions, the course of the moult cycle, biomass (dry weight, W; carbon, C; nitrogen, N; hydrogen, H; energy, E; the latter estimated from C), and ecdysteroid titers (measured with a radio-immuno-assay as ecdysone equivalents) were investigated. When the larvae reached the PRS, they had gained 90% in W, 72% in C, 32% in N, 53% in H, and 65% in E, since hatching, corresponding to an accumulation of 87% of final W and 62 to 69% of C, N, and H reached later, at the end of the mould cycle in the control. The period of starvation caused a 2.5-d delay of the moult cycle, mainly in late premoult, and significant losses of biomass and energy. Starved and fed larvae secreted similar amounts of moulting hormone per individual, but with a reduced rate in the starved group, thus causing developmental delay. Zoea II larvae moulting after starvation contained less than half of the control biomass and energy, and even less than a freshly hatched zoea I. Growth rate was only slightly enhanced in these zoea II larvae as compared to the fed control, but losses of biomass, mainly of lipids, were partly compensated by a 4-d prolongation of their moult cycle, chiefly (3 d) in stage C. Biomass curves were almost parallel in the two experimental groups of zoea II larvae, with significantly higher values in the control during all stages of the moult cycle. However, similar relative proportions (74 to 89%) of late premoult biomass and energy were reached at the D₀ threshold, regardless of different feeding history and initial or final values in a given group. The ecdysteroid titer curve of the zoea II which had moulted from starved zoea I was very similar to that in control larvae, but with a 3-d delay in the occurrence of premoult peak concentration (in both groups in stage D₁). Regulation and coordination of moult cycle, ecdysteroid titers, and growth in the larval development of decapod crustaceans are discussed, with special reference to the D₀ threshold.     

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.     

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.