The behavior of heterotypic resting schools of juvenile grunts (Pomadasyidae)

Resting schools of juvenile grunts (less than 12 cm length), composed primarily of Haemulon flavolineatum and H. plumieri, were studied from 1972 to 1976 on a series of patch reefs surrounded by seagrass beds on the northeast coast of St. Croix, U.S. Virgin Islands. Juvenile grunts form large inactive multispecies schools in reef areas by day. Repeated censuses over several years reveal that certain parts of the reef, particularly those with large formations of the corals Porites porites and Acropora palmata, are traditional schooling areas. When the fishes reach a size of about 15 cm, the H. plumieri form homotypic schools which roam the patch reefs but are no longer associated with particular formations of coral. In contrast, larger H. flavolineatum are uncommon on the patch reefs and they may move offshore to deeper water. The schools of juvenile grunts migrate some distance away (often 100 to 300 m) into seagrass-covered areas to feed on invertebrates at dusk each day. The routes which are followed into the seagrass beds are precise and persist over years. At dawn the grunts return to the reef on the same routes. Mass marking and release experiments show that juvenile grunts will move over long distances (approximately 3 km) to reach a home reef.     

Role of vision in the aggregative behavior of the planktonic mysid Mysidium columbiae

 Aggregations of the planktonic mysid Mysidium columbiae use vision to help maintain position within mangrove prop-root habitats and to maintain spacing within schools. Laboratory studies show that mysids videotaped in darkness using infrared illumination did not form schools and could not hold position in currents. In daylight, mysids more effectively held position in a flow-through chamber with a high-contrast visual reference than in its absence. The compound eyes of mysids are thought to be good motion detectors, but little is known about their visual acuity or sensitivity. An optokinetic drum was used to test the visual acuity and photosensitivity of mysids using the behavioral response of mysids to vertical black and white stripes that move past their field of view. When the drum rotates, the mysids swim at the same speed and in the same direction as the moving stripes. Swimming speed of the mysids was measured to compare their speed to the turning rate of the drum using video-computer motion-analysis techniques. Detection of the moving stripes was also inferred from the proportion of mysids that followed the stripes and that reversed direction when the rotation of the drum was reversed. By varying the width of these stripes, the visual acuity of the mysids was determined. The ability of M. columbiae to follow stripes of 1 mm in width from a distance of 15 to 30 mm indicates that mysids can visually resolve nearby prop roots and other mysids within schools. The photosensitivity threshold for the optokinetic response was found to be 0.001 μm photons m⁻² s⁻¹, similar to light levels during moonlight. These mysids are potential prey to a wide range of planktivorous fish, and their survival may depend upon their ability to maintain their position within schools and within the safety of the prop-root habitat during daylight hours in spite of currents and turbulence that would tend to disperse them. Received: 3 December 1999 / Accepted: 22 May 2000     

Growth and moulting of Neomysis integer (Crustacea: Mysidacea)

The growth and moulting of Neomysis integer (Leach) was investigated in the field and the laboratory. In the Ythan estuary, Aberdeenshire, Scotland, monthly samples taken from November 1976 to October 1978 revealed that the summer generation juveniles and mature individuals grew at a rate of 4 to 5 mm and 1 to 2 mm monthly, respectively. The winter generation had a growth rate of 3 to 4 mm monthly for juveniles and about 1 mm for mature individuals; during the winter there was a period of 3 mo when growth was almost completely stopped. Mysids reared in the laboratory on Artemia sp. nauplii had an average daily growth rate of 0.06 mm at 9°C and 0.09 mm at 16°C. The growth factors of N. integer ranged from 3 to 17% for mature and immature individuals, respectively. Intermoult periods ranged from 3 to 7 d in immature mysids to 12 to 18 d in mature mysids. Average laboratory growth curves calculated from information on growth factors and intermoult periods indicate that at 9°C (winter generation) it takes N. integer 277 d to grow to be a 15 mm mature individual, whereas at 16°C (summer generation) it takes 188 d. N. integer moults 24 times as it grows from a juvenile to a mature individual.     

Resident mysids: community structure,abundance and small-scale distributions in a coral reef lagoon

Seasonal and diel variations in community structure and abundance of coral-reef lagoon mysids were examined at Davies Reef in the central region of the Great Barrier Reef (GBR) between June 1980 and May 1981. Twenty-five mysid species belonging to three subfamilies of the family Mysidae were captured during the study, with six new records for the GBR. The epibenthic mysid community differed from that in the overlying water, was faunistically uniform, but formed characteristic seasonal and diel groupings. The dominant epibenthic species were Erythrops sp., Anisomysis pelewensis, Doxomysis littoralis, A. laticauda, Prionomysis stenolepis, A. lamellicauda, and A. australis, five of which formed schools. Total mysid abundances ranged between 110 and 790 m^-3 with peak abundance in October. Schooling species occurred at local densities of up to 500 000 m^-3. Mysids were absent from shallow and midwater depths during the day, but were distributed throughout all depths at night with peak abundances in mid-water and deep layers. The dominant species in the water column at night were Pseudanchialina inermis, A. laticauda and Gastrosaccus indicus, in descending order of abundance. Lagoonal mysids contribute little to the food of sessile reef planktivores, as all but three species remain concentrated near or on the lagoon floor both day and night. The contribution of resident lagoon mysids to reef trophodynamics is probably through remineralization of lagoon detritus. Given the vast reef areas comprised of sandy lagoons, the large populations and relatively large size of lagoon mysids, this trophodynamic role may be of considerable importance.A.I.M.S. Contribution No. 477     

Temporal patterns and transport processes in recruitment of spiny lobster (Panulirus argus) postlarvae to south Florida

We used time-series analysis to identify weekly and annual patterns in the supply of spiny lobster, Panulirus argus (Latreille, 1804), postlarvae to the Florida Keys, USA, over an 8 yr period. We also investigated the relationship between postlarval influx and wind forcing as a transport mechanism using the complex vector-scalar correlation analysis. Postlarval supply had a lunar phase periodicity at 4.5 wk intervals, with postlarval abundance peaking between the new moon and first-quarter lunar phases. A distinct annual cycle of postlarval supply with two peak periods was also apparent. Cross-correlation analysis between relative postlarval abundance and a 12 mo cycle showed that the annual peak occurs in spring, centered around March. With the 12 mo periodicity removed, a smaller peak at 5 mo intervals was also well defined. Wind-forcing for 7 d prior to the time of postlarval collection was marginally correlated with postlarval abundance through the entire time-series; the association was strongest during the late fall to early spring months. The analysis indicated that postlarval supply was correlated with winds from the northeast (ca. 45°), which are associated with winter atmospheric fronts. In contrast to results reported for other spiny lobster populations, these patterns suggest that recruitment of lobster postlarvae to south Florida is predictable only at a gross level and is presumably affected by the temporally inconsistent structure of regional oceanic gyres and variability in the timing of lobster spawning in the Caribbean. Received: 24 January 1997 / Accepted: 4 March 1997     

Food utilisation of pelagic mysids, Mysis mixta and M. relicta, during their growing season in the northern Baltic Sea

The dietary habits of the pelagic mysid Mysismixta were studied during its growing season at an open sea location in the Gulf of Finland, the northern Baltic Sea. Stomach samples were taken twice a month from June to September 1997. The most abundant phytoplankton taxa in the stomachs were diatoms and dinoflagellates, and copepods and cladocerans were the most abundant zooplankton identified. A clear change was found in the diets during the study period. Small mysids (3 to 6 mm) fed on sedimented phytoplankton in the early summer (90% benthic particles in June) but shifted gradually to a more pelagic and carnivorous diet (>40% pelagic particles, consisting of ca. 60% zooplankton in September). Seasonal changes in mysid capture ability as well as food availability were suggested to affect the diet composition of mysids during their growth. The ratio of pelagic and benthic food particles could – irrespective of the season – be explained by mysid size, whereas the zooplankton:phytoplankton ratio was better explained by season. The stomach analysis suggests that the mysids needed to attain a threshold size of 8 to 11 mm to initiate feeding on the more evasive copepods. Mysids also started to grow faster at the same time as the proportion of copepods increased in the diet, which suggests that copepods are an important energy source for M. mixta in late summer. Finally, a comparison was made between the M. mixta diet and that of the less abundant M. relicta. The diets of the two pelagic mysid species overlapped by 75% (Schoener's index). The main difference was due to M. mixta eating more zooplankton and pelagic material than M. relicta. Received: 15 September 1999 / Accepted: 18 January 2000     

Cercopagis pengoi and Mysis spp. alter their feeding rate and prey selection under predation risk of herring (Clupea harengus membras)

The anti-predator behaviour of Baltic crustacean planktivores was studied in feeding experiments under predation pressure of herring. The experiments were conducted with pelagic mysids: Mysis mixta and Mysis relicta, and with Cercopagis pengoi, a non-indigenous cladoceran, which invaded the Baltic Sea in 1992. Zooplankton was offered as prey. Two kinds of experiments were performed in the absence and presence of chemical predator cues: (1) two-prey experiments with prey, which have poor or good escape responses and all three planktivores and (2) natural prey experiments with mysids in natural zooplankton assemblages. The results showed that all three species reacted to the chemical cue of herring by decreasing their feeding rate and altering prey selection. C. pengoi selected easily captured prey (rotifers) in two-prey experiments under predation risk while selection for any prey was evident in mysids in natural prey experiments only in the absence of predator cues. This indicates that planktivores have different anti-predator strategies, which are modified by their own prey capture abilities. C. pengoi was a very efficient predator on small prey with size-specific prey consumption rate 5 to 18 times the rate of mysids. Results show that the studied planktivores are capable of adjusting their feeding behaviour to decrease their conspicuousness in order to increase survival under predation risk. Further, results support the view that C. pengoi has adapted well to the Baltic ecosystem, sharing food niche with pelagic mysids and most probably having a strong influence on the whole pelagic food web.     

Larval development in the Tasmanian mysids Anisomysis mixta australis, Tenagomysis tasmaniae and Paramesopodopsis rufa (Crustacea: Mysidacea)

We describe the brood duration and marsupial development of three temperate coastal mysid species, Anisomysis mixta australis (Zimmer), Tenagomysis tasmaniae Fenton and Paramesopodopsis rufa Fenton, found commonly in Tasmanian waters. Larvae cultured in vitro had brood durations at 13 °C (17 °C) of 22 (15), 23 (15), and 28 (20) d, respectively. Development through seven larval stages, and brood durations for these three species are similar to those reported for coastal mysids from other temperate areas throughout the world. Received: 10 June 1997 / Accepted: 1 July 1997     

Effect of mercury on the survival,respiration, and growth of postlarval white shrimp,Penaeus setiferus

The results of toxicity experiments have shown that a level of 17 ppb ionic mercury is acutely toxic (96-h LC₅₀) to postlarval white shrimp (Penaeus setiferus). In addition, the size of the shrimp, within the range tested (7 to 35 mm) did not significantly alter the toxicity values of mercury to this species. Fifty-seven days pre-exposure to low levels of mercury (0.5 and 1.0 ppb) had no effect on the acute toxicity (96-h LC₅₀) of mercury to the shrimp. The effect of chronic exposure to low levels of mercury on the respiratory rate, growth, and molting rate of postlarval P. setiferus was studied, and the results showed that 60 days exposure to 0.5 and 1.0 ppb mercury did not significantly affect any of these parameters for the postlarvae.     

Turbinaria ornata as an herbivory refuge for associate algae

Habitat associations are an integral part of coral reef community structure. Commonly, one organism lives in such close association within or near another that a spatial refuge occurs, whereby one of the organisms provides protection to the other. This is often the result of defenses of the host deterring an associate organism’s consumers. In Moorea, French Polynesia, the range and abundance of the brown macroalga, Turbinaria ornata, have increased drastically since 1980 such that dense aggregations of this macroalga are a dominant component of the backreef habitat. Turbinaria ornata is both mechanically and chemically defended from herbivores. Other species of macroalgae grow within aggregations of Turbinaria and may benefit from these defenses. This study investigates whether aggregations of Turbinaria create a refuge from herbivory for associate macroalgae. When Turbinaria aggregations were removed experimentally, there was a significant increase in the number of associate algal species. Moreover, an herbivory assay using the palatable local alga Acanthophora spicifera identified herbivory as the mechanism for lower diversity on bommies lacking Turbinaria aggregations. The local increase in algal richness due to the refuge from herbivory afforded by Turbinaria may be an important contribution to macroalgal and community dynamics on reefs in Moorea, French Polynesia.     

Effects of various diets on the nutritional value of Artemia sp. as food for the prawn Penaeus monodon

The nutritional value of Artemia sp. as food for marine fish and crustacean larvae has been linked to the level of its polyunsaturated fatty acid (PUFA) content. Experiments in August 1984 were conducted to determine the effects of various artificial diets and algae on fatty acid composition of PUFA-deficient Artemia sp. (Utah GSL strain) and their resulting value as food for postlarvae of the prawn Penaeus monodon (Fabricius). Nauplii of the brine shrimp were grown on extracts of corn, copra, soybean and rice bran containing precursors (C18) to long-chain PUFA and also on algal species containing different levels of long-chain PUFA (C20). The nauplii were then used as food for P. monodon postlarvae. The results revealed that absence of C20 polyunsaturates from the feeds and their presence in the algae were reflected in the polyunsaturated fatty acid content of the tissues of Artemia sp. When fed with brine shrimp fed on algae, P. monodon displayed better postlarval survival and significantly higher growth; related to the content of polyunsaturated fatty acids in Artemia sp. A practical feeding approach in prawn hatcheries would be to grow Artemia sp. on a cheap diet such as rice bran, and then to enhance its nutritional value with a diet high in PUFA prior to harvesting, in order to improve hatchery production.     

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     

Changes in nucleic acids over the molt cycle in relation to food availability and temperature in Homarus americanus postlarvae

Postlarval lobsters Homarus americanus Milne Edwards hatched from three females collected in 1989 from Block Island Sound, Rhode Island were reared individually in the laboratory under nine treatment combinations of temperature (15, 18 and 21°C) and feeding (starved, low ration and full ration). Total RNA, DNA (mgind.^-1), RNA:DNA ratios and molt stage were determined for individuals sampled at daily intervals. Postlarval lobsters had high resistance to starvation. A majority of the lobsters survive 12 d of food deprivation, with some surviving up to 24–29 d. During starvation, cell biomass (estimated from protein:DNA) decreased to a minumum size, whereas cell number (based on total DNA) was generally conserved. The molt cycle was arrested at molt stage C in the starved postalrvae. Instar duration was inversely related to temperature. However, the duration of the postlarval instar did not differ between the low and full ration treatments. Uncoupling of cell growth and the molt cycle was evident in the full and low ration treatments. In the full ration treatments, the postmolt through early premolt was characterized by a rapid increase in total RNA and DNA. Maximum cell biomass was attained by molt stage C or D₀. In the low ration treatments, total RNA and DNA were less than those in the full ration treatments and the maximum cell biomass was attained only at molt stage D_2-3. Notably, different feeding regimes resulted in different patterns in RNA:DNA ratios over the molt cycle during the postlarval instar. RNA:DNA ratio was a sensitive indicator of recent (2 to 3 d) food deprivation. However, this ratio was positively related to the level of feeding only at molt stages C to D₁ and was inversely related to temperature regime. These results indicate that the use of RNA:DNA ratios to estimate the relative nutritional state of postlarval tobsters must be qualified with respect to the period of the molt cycle and the temperature regime.     

Influence of delayed metamorphosis on survival and growth of juvenile barnacles Balanus amphitrite

Competent cyprid larvae of the barnacle Balanus amphitrite Darwin were prevented from metamorphosing in the laboratory for 3 or 5 d using three different techniques (holding at low temperature, crowding, and detaining on a silanized surface). We then assessed the effects of prolonging larval life on post-metamorphic growth and survival, in comparison with control individuals that metamorphosed soon after they were competent to do so. Seven experiments were conducted over 2 yr (July 1987 to September 1989). In all experiments (each with six replicates per treatment), postponing larval metamorphosis for 3 or 5 d dramatically depressed postmetamorphic growth rate (P<0.05), although metamorphic success and post-metamorphic survival were not affected (P>0.10). The results suggest that B. amphitrite cyprids deferring their metamorphosis in the field may be less successful in competing for space, at least during the first few weeks of postlarval life.     

Life histories of three species of lanternfishes (Pisces: Myctophidae) from the eastern Gulf of Mexico

Morphology and microstructure of the sagittal otoliths from three species of mesopelagic, tropical-subtropical myctophids [Benthosema suborbitale (Gilbert),Diaphus dumerilii (Bleeker)Lepidophanes guentheri (Goode and Bean)], collected from September 1984 to May 1986 in the eastern Gulf of Mexico (27°N, 86°W), were examined and described. Analysis of the microstructure revealed microincrements corresponding to the daily growth rings reported in many studies. Using marginal increment analysis, the deposition of microincrements was verified as occurring daily, the first validation of daily growth rings in the otoliths of mesopelagic fishes. In all three species, the clear central (larval growth) zone of the sagitta was sharply delimited by a dark check accompanied by a series of accessory primordia. A wide, dark, sharply defined postlarval zone (width 100 µm) radiated out from this boundary inB. suborbitale andL. guentheri. While a darkened region was also observed around the larval growth zone inD. dumerilii, it was diffuse and differed in structure from that in the other two species. Within the dark zone in the otoliths ofB. suborbitale andL. guentheri, two different microincremental structures were observed. The narrower of these was determined to be the increment deposited on a daily basis. The structure and formation of the dark region in these three species appear to be related to larval transformation and behaviors of different species of myctophid larvae.     

Predatory feeding of two marine mysids

Predatory feeding of the marine mysids Mysidopsis bigelowi and Neomysis americana on several species of co-occurring copepods was examined in laboratory experiments. M. bigelowi exhibited a curvilinear functional response; there was a negative logarithmic relationship between prey density and clearance rates. N. americana also exhibited higher clearance rates at lower prey densities. Increased clearance rates at lower prey densities were probably due to increased swimming speed or reaction distance as hunger increased. This response occurred only when mysids could visually locate prey; in complete darkness clearance rates were significantly lower and independent of prey density. Feeding rates on different prey species were only partially dependent on prey size; prey movement patterns and escape behavior also strongly affected feeding rates. M. bigelowi showed active prey selection when offered a choice of different prey species. Estimates of predation rates of estuarine mysid populations indicate that they could have a significant effect on co-occurring copepod populations.     

Predation of a sand-dwelling mysid crustacean Gastrosaccus sanctus by plover birds (Charadriidae)

The plovers Charadrius hiaticula and C. alexandrinus feed on the mysid shrimp Gastrosaccus sanctus (van Beneden, 1861). They hunt at random along the sandy seashore, on the sites of the densest mysid population, regardless of the size of the mysids.     

Behaviour and tolerance to hypoxia in juvenile Norway lobster (Nephrops norvegicus ) of different ages

The annual occurrence of hypoxia (<25% oxygen saturation) in the bottom waters along the Swedish west coast coincides with the postlarval settlement of Norway lobster, Nephrops norvegicus (L.). This study investigates behaviour and the experimental effects of low oxygen concentrations in juvenile N. norvegicus of different ages. All experimental individuals were reared to the juvenile (postlarval) stage in the laboratory and then given sediment as a substratum. Behavioural responses to low oxygen concentrations were tested in early and late Postlarvae 1 exposed to normoxia (>80% oxygen saturation, pO₂ > 16.7 kPa), moderate hypoxia (30% oxygen saturation, pO₂ = 6.3 kPa) and hypoxia (25% oxygen saturation, pO₂ = 5.2 kPa). The experiments were run for a maximum period of 24 h or until individuals died. Behaviour was studied using sequential video recordings of four behavioural activities: digging, walking, inactivity or flight (escape swimming up into the water column). Behaviour and mortality changed with lowered oxygen concentrations; energetically costly activities (such as walking) were reduced, and activity in general declined. In normoxia, juveniles initially walked and then burrowed, but when exposed to hypoxia they were mainly inactive with occasional outbursts of escape swimming. To increase oxygen availability the juveniles were observed to raise their bodies on stilted legs (similar to adults in hypoxic conditions), but oxygen saturations of 25% were lethal within 24 h. The results suggest that the main gas exchanges of early postlarval stages occur over the general body surface. Burrowing behaviour was tested in Postlarvae 1 and 2 of different ages held in >80% oxygen saturation for 1 wk. The difference in time taken to complete a V-shaped depression or a U-shaped burrow was measured. The results showed a strong negative relationship between postlarval age and burrowing time, but all individuals made a burrow. Juveniles were more sensitive to hypoxia than adults. Thus, the possible consequences of episodic hypoxia for the recruitment of Nephrops norvegicus and for the recolonization of severely affected areas are discussed. Received: 4 August 1996 / Accepted: 11 October 1996     

Potentiometric studies of mixed complexes of cobalt(II) and copper(II) with l‐Asparagine and adenine

A potentiometric titration technique has been used to determine the stability constants for the various complexes of Co(II) and Cu(II) with L‐asparagine and from DNA base, e.g. adenine. Stability constants of ternary systems have been evaluated by the method suggested by Irwing‐Rossotti. In addition, the conditional constants were calculated as a function of pH. The maximum values of the conditional formation constants were found to be in accordance with the mixed‐ligand complex formation constants in a determined pH region. Furthermore, the molar fractions of different species from mixed complexes were calculated by means of formation constants. The values of stability constants of mixed‐ligand complexes at 25°C are as follows: log K= 5.25 for Co(II)‐L‐asparagine‐adenine; log K= 9.30 for Cu(II)‐L‐asparagine‐adenine. The ionic strength was kept constant at I = 0.20 with NaClO₄.     

Ontogenetic variation in digestive proteinase activity of larvae and postlarvae of the pink shrimp Farfantepenaeus paulensis (Crustacea: Decapoda: Penaeidae)

Proteinase (endopeptidase), trypsin-like and chymotrypsin-like activities were examined throughout the ontogenetic development of cultured Farfantepenaeus paulensis. Whole individuals from different larval and postlarval stages, and the hepatopancreas of adults were homogenized and assayed to quantify the enzyme activities of specific substrates. Proteinase activity was identified by substrate-SDS-polyacrylamide gel electrophoresis. Specific inhibitors for trypsin (TLCK), chymotrypsin (TPCK) and serine proteinases (PMSF) were used to identify activity zones of these enzymes in gels. Protein-specific activity of total proteinases, trypsin and chymotrypsin was negligible at the egg stage and at Nauplius III, increasing in the first protozoeal substage (PZ I), and reaching a peak at PZ III; it decreased again in the subsequent postlarval substages. Different patterns of proteinase activity were observed in SDS-PAGE zymograms during ontogenetic development. Active bands of 14.6, 16.4, 17.5, 19.5, 22.5, 23.9, 25.8, 28.9, 32.0, 34.4, 37.7, and 42.2 kdaltons were detected in the adult hepatopancreas. Proteolytic activity was detected on gels in PZ I, and intense activity zones of 16.4, 17.5 and 19.5 kdaltons were found up to Mysis I (M I). Intense bands of 39.1 and 53.5 kdaltons were observed only at PZ III and M I. Band-activity intensity decreased after metamorphosis to the postlarval stage (PL). The chymotrypsin inhibitor TPCK had no effect on the proteinase bands. Active zones in gel inhibited with both TLCK and PMSF were considered to represent trypsin. The inhibitory effect of PMSF alone on proteinase extracts indicated chymotrypsin activity. TLCK and PMSF inhibition also varied during ontogenetic development. The inhibition of bands recorded between 14.6 and 21.7 kdaltons suggested the presence of low molecular weight trypsin in F. paulensis. The 39.1 kdaltons band observed at PZ III and M I were trypsin-like. On the other hand, bands of 28.9, 32 and 37.7 kdaltons from the adult hepatopancreas seem to represent a chymotrypsin. We conclude that the recorded variation in enzyme activity may be associated with morphological and behavioral changes during penaeid ontogenetic development. The higher enzyme activity at PZ II, PZ III and M I may reflect the increased energy turnover associated with intense swimming behavior and food ingestion. Received: 24 September 1998 / Accepted: 20 August 1999