Managing the offspring environment: brood care in the bromeliad crab,Metopaulias depressus

Summary Maternal modification of the larval environment was studied in the bromeliad crab, Metopaulias depressus (Decapoda, Grapsidae). The bromeliad crab breeds in rain water stored in the leaf axils of large Jamaican bromeliads. The physico-chemical water conditions are highly unfavorable for the development of the larvae. Water in nursery axils contained no leaf litter and up to 14 empty snail shells, whereas other axils were partly or completely covered with leaf litter and rarely held snail shells. Mother crabs removed leaf litter from nursery axils, and occasionally from neighboring axils. Nursery axils contained twice as much dissolved oxygen (DO₂) at night when DO₂ is at a minimum, and less dissolved carbon dioxide (DCO₂) than axils unattended by a crab. The higher DO₂ content in nurseries meets the respiratory needs of an average sized brood, but the lower concentration in axils with litter may not. It is suggested that the higher DO₂ and lower DCO₂ concentrations in the nursery axils result from a water circulation produced by the mother and her leaf litter cleaning behavior. The low pH and the Ca²⁺ content in the water of unattended bromeliad axils are unfavorable for development of larvae and young. Mother crabs significantly increased pH and Ca²⁺ by adding empty snail shells to the nursery-axil water. In experiments where the brood was transferred to other axils, mothers switched from the nursery axil to the new axil containing the brood, and there continued caring for their offspring.     

Developmental changes in body weight and elemental composition of laboratory-reared larvae of the red frog crab,Ranina ranina (Decapoda: Brachyura)

Ranina ranina larvae were reared at 29°C from hatching to the megalopa stage to measure daily changes in body weight, water content and elemental composition. Energy, estimated from carbon content, was also examined. The water content was 85 to 92% of body weight immediately after ecdysis but decreased with days after ecdysis. Gains in body weight, carbon, nitrogen, hydrogen and energy during each instar ranged from 52 to 245% and increased with instar after instar II (body weight and carbon), instar III (hydrogen and energy), and instar IV (nitrogen). Cumulative gains of these elements from hatching to 2 d before metamorphosis into megalopa ranged from 11 567% (carbon) to 12 209% (energy). Most cumulative gains (57 to 59%) in elemental composition were contributed by instar VII. Carbon, nitrogen and hydrogen content in body weight decreased to a minimum on the day of ecdysis and increased on the subsequent days. C:N ratios after instar IV were lowest on the day after ecdysis and reached a plateau by the second day. Energy, estimated as J mg^-1 dry weight (DW), decreased with instar and within a molt cycle, and was at a minimum on the day after ecdysis. Gains in elemental composition could be described by an exponential function of days after hatching and by a quadratic function in each instar.     

Vertical migrations in the tree crab Sesarma leptosoma (Decapoda,Grapsidae)

Sesarma leptosoma, somewhat similar to the Atlantic related species, Aratus pisonii, is an East-African mangrove crab which spends its entire life on the roots and branches of mangrove trees (mostly Rhizophora mucronata, Bruguiera gymnorhiza and Ceriops tagal). S. leptosoma never enters the water, nor does it ever venture onto the free mud surface at low tide. Part of the day and night it remains on the lower parts of the mangrove aerial roots, which are often encrusted with a wet algal mat of Bostrichia spp., searching for food and water. Twice a day, from ca. 06:00 to 08:00 hrs in the morning and 16:00 to 18:00 hrs in the afternoon, many of the crabs migrate as far as the leaves at the top of the tree on which they feed. However, they only spend a brief period among the leaves, from ca. 07:00 to 10:00 and 17:00 to 19:00 hrs, after which time they make their way back towards the roots again in two downward migrations. In the morning, the downward migration brings all the crabs back to the roots, but in the evening not all the crabs take part in the downwards migration and some of them pass the whole night in the tree tops. A comparison of the migration time patterns for two different periods of the year (June–July and November) shows that the number of crabs migrating along the tree trunk is modulated by the spring-neap tidal cycle, while the onset of daily migration seems to be controlled mostly by the light level and/or other climatological cues. The adaptive significance of this migratory behaviour is discussed. Observations reported in our study were made in Mida Creek, Dabaso, Kenya in 1991 and 1992.     

Timing of copulation in the molting and reproductive cycles in a grapsid crab,Gaetice depressus (Crustacea: Brachyura)

From the seasonal changes in the amount of semen in the sperm ducts of males and in the fullness of spermatheca of females, the copulation season of Gaetice depressus in central Japan is considered to approximately coincide with the oviposition season, from March to October. The sperm ducts of all the large males examined contained semen, the amount of which did not vary obviously with stages of the molt cycle or with seasons. The genital apertures of females were normally covered with calcified immobile opercula in this species. The opercula decalcified after molting and again shortly before oviposition, and retained mobility for 1 to 3 d. Shortly after molting, soft-shelled females with mobile opercula did not have well-developed ovaries nor did they copulate with males even if they had a chance. In contrast, all of the hard-shelled females with mobile opercula had well-developed ovaries, and laid eggs within 2 d after the decalcification of opercula regardless of the occurrence of copulation. Laboratory experiments showed that the females normally copulated with males before oviposition. Crabs examined in this study were collected from Shirahama, Wakayama Prefecture, Japan between March 1988 and March 1989.     

Acute temperature and low dissolved oxygen tolerances of brachyuran crab (Cancer irroratus) larvae

Tolerance limits to acute temperature, and temperature and low dissolved oxygen stresses were determined for 5 zoeal stages and megalops of Cancer irroratus Say. The acute temperature limits for a 120 min exposure were all approximately 29.0°C, with little interstage, variation, while those for 240 min exposure ranged from 27.3° to 28.5°C. More interstage variation was shown when temperature and low dissolved oxygen stress were combined, with low oxygen tolerance decreasing as temperature increased. The first, second, and fourth zoeal stages display similar patterns of response. The third and fifth stages show a similar response, but are different from the former stages. The megalops is relatively insensitive to changes in oxygen concentration with temperature. The larval stages did not show a progressive increase in tolerance to temperature or low dissolved oxygen with development. Tolerance to these factors may be related to size, stage of development and activity level. The larval stages have capacities to tolerate a wider range of these factors than they require in the natural environment.     

Rhythms in larval release of the sublittoral crab Neopanope sayi and the supralittoral crab Sesarma cinereum (Decapoda: Brachyura)

Ovigerous females of the subtidal xanthid crab Neopanope sayi (Smith) and the high intertidal grapsid crab Sesarma cinereum (Bosc) were collected during the summers of 1986 and 1987 in the Beaufort, North Carolina (USA), area and brought into the laboratory, where rhythms in larval release were monitored. When crabs with late-stage embryos were put under a 14 h light:10 h dark cycle in an otherwise constant-environment room, an apparent tidal rhythm in release of larvae was observed for both species, with N. sayi releasing near the time of day and night high tides, and S. cinereum releasing around the time of night high tides. The time of sunset relative to high tide was a complicating factor, since larval release for both species was often concentrated around sunset when evening high tides fell several hours before sunset. When a group of N. sayi and S. cinereum were brought into the laboratory and placed under constant lowlevel light for 5 d, the release rhythm of the population persisted, thus implying that the rhythm is endogenous. Larval release near the time of high tide and often at night is common among brachyurans living in tidal areas, regardless of specific adult habitat, suggesting a common functional advantage. Possibilities include transport of larvae from areas where predation and the likelihood of stranding and exposure to low-salinity waters are high, as well as a reduced probability of predation on adult females. Results of the present study suggest that the importance of release after darkness may increase with increasing tidal height of the adult.     

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.     

Population dynamics of the commensal spider crab Inachus phalangium (Decapoda: Maiidae)

The ecology of the spider crab Inachus phalangium (Fabricius, 1775) (Decapoda: Maiidae) was studied in the field. I. phalangium inhabits the sublittoral on the sea anemone Anemonia sulcata Pennant. From July 1981 to April 1984 in the Mediterranean (Banyuls sur Mer, southern France) more than 3000 anemones were examined and ca 1800 I. phalangium were found on them. The population dynamics' generation cycles, reproductive activities and the dynamics of the sex ratio were investigated. The density of juveniles (crabs before the pubertal moult) on anemones changed in a yearly cycle from low in the first six months to very high in the second six months. The first occasional young crabs of a generation appeared in March/April (3rd and 4th decapodite stages) on the anemones. Their density increased enormously in the following months. The generation grew gradually on the anemones and moulted into puberty in September-January. Density of adults (crabs after the pubertal moult) on anemones changed in a yearly cycle from low to high from the summer to winter months. A new adult generation was recruited every autumn through the pubertal moult and disappeared in the following summer. Female reproductive activity continued throughout the year. Females carried several broods in succession, but the frequency of breeding females fluctuated on a yearly cycle. The highest percentage of egg-carrying females, i.e. the peak of the reproductive season, lay in the first half of the year. The maximal life span of a generation, from the hatching of the first larvae to the disappearance of the last adults, lasted 1.5 to 2 years. Males moulted into puberty ca one month later than females. The moulting distribution of adult males had roughly the same course as in females. An adult male generation diet out about one to two months before the female generation. Life expectancy was therefore 14 to 17 months for females and 12 to 15 months for males. The sex ratio of juveniles shortly before the pubertal moult was balanced. The sex ratio of adults shifted from 1:1 at the beginning of the reproductive period to ca 1:9 in favour of females at its end.     

Growth and elemental composition (C,N, H) in Inachus dorsettensis (Decapoda: Majidae) larvae reared in the laboratory

Larvae of the spider crab Inachus dorsettensis were reared in the laboratory at constant 12 °C. Development lasted 8 to 10 d in the Zoea I, 10 to 12 d in the Zoea II and 14 to 20 d in the megalopa stage. During this time, larval growth was measured in samples taken every 2 to 4 d as dry wt (W), carbon (C), nitrogen (N), and hydrogen (H); energy content (E) was calculated from C. Biomass and energy (per individual) increased in each larval stage as a parabola-shaped function of age, which could be fitted by a power equation. C, H, and E show a higher percentage gain (relative to the initial values at hatching) than W or N, suggesting that proportionally more lipid than protein is accumulated during larval development. There are cyclical changes in the relative (per unit of W) biomass and energy figures, corresponding to the larval moult cycles: immediately after each ecdysis all these values decrease, presumably due to rapid uptake of water and minerals, then they increase again due to tissue growth and remain high until the next moulting occurs. Cyclical changes in the C/N ratio suggest that proportionally more lipid than protein is accumulated during the initial (postmoult) phase of the moult cycle, followed by a period of balanced or protein-dominated gain during the intermoult and premoult phases. These patterns of growth and elemental composition observed during the complete larval development and in single moult cycles of I. dorsettensis are compared with those described in the literature for other decapod species. This comparison suggests a high degree of similarity in biochemical composition and growth characteristics of larval decapod crustaceans.     

Preliminary rearing experiments on the larvae of Sergestes lucens (Penaedia,Natantia, Decapoda)

Sergestes lucens Hansen, a mesopelagic shrimp fished commercially in Suruga Bay, Japan, was successfully reared from egg to post-larval stage V under laboratory conditions. Chaetoceros ceratosporum and Artemia nauplii were found to be satisfactory food in the laboratory during rearing. Growth, mortality, food preference, and feeding and swimming activities during the various developmental stages were investigated. Temperature changes greatly affected the speed of development and the mortality of the larvae. The optimum temperature range for larval development was 18° to 25°C. The growth rate (length) of larval stages was as rapid as 0.16mm/ day at 20 °C and 0.21 mm/day at 23 °C. The larvae first started feeding on phytoplankton at elaphocaris stage I, and then gradually became predators in the post-larval stages. It is suggested that the critical period for the species occurs in the elaphocaris stages. Environmental data, vertical distribution of the species, and data obtained from laboratory experiments suggest that the fluctuation in the abundance of S. lucens is greatly influenced by the water temperature at around 50 m from June to August. Feeding mechanisms observed in the post-larval stages are described.     

Larval development of the spider crab Pisoides edwardsi (Decapoda,Brachyura) under laboratory conditions

Larvae of Pisoides edwardsi (Bell, 1835) have been reared in the laboratory at 2 different temperatures (13.8° and 18.5°C), from hatching to megalops stage. The two zoeal stages and the megalops, as well as the setation of the functional appendages are described and illustrated. The main characteristics useful to differentiate the larvae of P. edwardsi from those of Libidoclaea granaria, the other Chilean species belonging to the same sub-family, are discussed. Data on duration of zoeal development, length of moulting intervals, and mortality at the 2 test temperatures, are also given.This study was financially supported, in part, by the Chilean Ministry of Agriculture.     

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.     

Population dynamics and growth of juveniles of the velvet swimming crab Necora puber (Decapoda: Portunidae)

Recruitment variability plays a critical role in determining local population densities of benthic organisms, but extreme vulnerability at the onset of juvenile life is a trait that is largely responsible for population survivorship trends. The aim of the present study was to determine the role of juvenile recruitment in the population structure of Necora puber. Juveniles of N. puber were collected from the lower intertidal of rocky shores of Plymouth Sound (southwest coast of the UK) and monthly size–frequency distribution were used to determine the dynamics and the growth of the population. The parameters of the von Bertalanffy growth function were estimated (K=0.281 year⁻¹; t ₀=0.043; C=0.103; and t _s=0.268) assuming a L _∞=105 mm. Growth was markedly seasonal and present results indicated a slower juvenile growth rate than described previously for N. puber. The recruitment period was extensive and was two times higher in 2001 than in 2000 at the start of the 1+ year, but levelled off at the end of the 1+ year class on three of the four shores studied. Instantaneous mortality as high as 5.1 year⁻¹(99.4% year⁻¹) was observed during the higher recruitment year. Early juvenile mortality appears to be density dependent and a demographic bottleneck appears to limit the number of juveniles on some shores.     

Larval development of the red crab Pleuroncodes monodon (Decapoda Anomura: Galatheidae) under laboratory conditions

Larvae of Pleuroncodes monodon (Milne-Edwards, 1837), a red crab of commercial importance in South America, were reared in the laboratory at 2 different temperatures (15° and 20°C), from hatching up to the last larval stage. The 5 typical stages, with their corresponding functional appendages, are described and figured. The main characteristics useful in differentiating larvae of P. monodon from those of the other Chilean species of Galatheidae and its northern congener P. planipes are discussed. Data on duration of each larval stage, length of moulting intervals and mortality at the 2 test temperatures are also given.This study was financially supported by the Chilean Ministry of Agriculture and by the National Commission for Scientific and Technological Research (CONICYT).     

Ultrastructural changes in the gill epithelium of the crab Chasmagnathus granulatus (Decapoda: Grapsidae) in diluted and concentrated seawater

Chasmagnathus granulatus Dana, 1851 is an intertidal estuarine crab that experiences acute salinity changes ranging from <1‰ to full-strength seawater and even hypersaline waters in tide pools concentrated by evaporation. Ultrastructural changes induced by salinity in the posterior gills were examined in crabs collected from the Rio de la Plata estuary Argentina during March 1999. The posterior gills of C. granulatus are involved both in ion uptake and ion secretion depending on the acclimation medium. These organs are mostly lined with a thick tissue, which presents the characteristics of a typical salt-transporting epithelium. Electron microscopy analysis of gill tissue from crabs acclimated to dilute, full, and concentrated seawater (12‰, 34‰, and 45‰ salinity) showed significant development of basolateral membrane interdigitations, with numerous mitochondria and conspicuous apical membrane infoldings. Morphometrical analysis indicated that the subcuticular space delimited by the infolding of the apical membrane was significantly increased in the gills of high-salinity acclimated crabs. Septate junctions, which are thought to define the paracellular permeability, were significantly shorter in high-salinity acclimated crabs, suggesting a possible role of the paracellular pathway in salt secretion.     

Sand disposal of the painted ghost crab Ocypode gaudichaudii (Decapoda: Ocypodidae): A possible role in courtship

In 1991/1992, we studied the sand disposal behavior of the painted ghost crabs Ocypode gaudichaudii on the Pacific coast of Panamá. O. gaudichaudii either kick, dump or tamp sand they excavate from their burrows. Here we relate these three kinds of sand disposal to burrow structure and distribution, as well as to crab size and sex. Our objective was to determine whether tamping may be a male courtship signal. Burrows whose owners tamped sand were on average longer, deeper, and higher on the beach than were burrows whose owners kicked or dumped sand. Five burrow shapes were distinguished, with half-spiral and spiral shapes being most common among tamped burrows. All crabs excavated from tamped burrows were males. Tamped burrows peaked in abundance around full and new moons. These observations, together with what is known about mating and breeding behavior of other ghost crabs, suggest that tamping may be involved in O. gaudichaudii courtship.     

On the introduction of the Kamchatka King crab Paralithodes camtschatica (Decapoda: Anomura: Lithodidae) into the Barents Sea

About 2,500 specimens of the Kamchatka king crab Paralithodes camtschatica (Tilesius), ca. 5 to 15 years old, and about 10,000 juveniles, ca. 1 to 3 years old, were released into the Barents Sea from 1961 to 1969. One large berried female was caught in 1974, and the number of records increased in 1976 (at least 100 records, including 6 reliable ones). The crab therefore seems to have become a permanent inhabitant of the Barents Sea. It will be important in future to continually record new occurrences of the crabs, especially of juveniles, in order to document the extent of their distributional range and to obtain evidence of successful reproduction of the species.     

Biochemical evidence for absence of feeding in puerulus larvae of the Western rock lobster Panulirus cygnus (Decapoda: Palinuridae)

Changes in energy-reserves during the transition from phyllosoma through peurulus to juvenile in the Western rock lobster Panulirus cygnus (George, 1962) were studied by means of carbon: nitrogen analysis. Specimens were collected by means of plankton nets and puerulus collectors along the Western Australian coastline between July 1992 and January 1993. Reserves are accumulated during the last phyllosoma stage and are consumed during the puerulus stage. These observations support the hypothesis that the puerulus is a non-feeding stage. Based on basal metabolic rate and observed changes in reserves, the natant puerulus stage is calculated to last a maximum of 21.6d. When the estimated cost of swimming is taken into account, the duration is reduced to perhaps a week. The latter time-span corre-sponds with field observations that both abundance of planktonic pueruli in oceanic waters and inshore settlement of pueruli peak in the week around new moon. At 18°C, the puerulus metamorphoses to the first juvenile stage within 12.8 d after settlement; at 23°C this is reduced to 8.3 d. After settlement, puerulus larvae have an energetic advantage at elevated temperatures; the increased cost of metabolism is compensated by an accelerated development. The natant puerulus, however, appears to be slightly disadvantaged energetically at elevated temperatures; both C:N ratio and ash-free dry weight of pueruli just after settlement show a decreasing trend as the water temperature increases. The effect of water temperature and the distance from the edge of the continental shelf to the shore on consumption of energy reserves might provide a mechanism to partially explain temporal and spatial patterns in puerulus settlement.     

Latitudinal variation in the reproductive biology of the commensal crab Pinnaxodes chilensis (Decapoda: Pinnotheridae) along the Chilean coast

The pinnotherid crab Pinnaxodes chilensis is a common commensal of the edible sea urchin Loxechinus albus along the Chilean coast. Several aspects of the reproductive biology of P. chilensis were examined between April and June 1999, along temperature and salinity gradients, at three sampling sites along the Chilean coast (23°45'S-39°24'S). Results demonstrated significant differences in egg number, egg volume, dry egg weight, and reproductive output of ovigerous females, between the studied populations of northern and central-southern Chile. Egg volume, egg dry weight, and reproductive output of females decreased from high to low latitudes, while egg number increased from high to low latitudes, exhibiting a clear trade-off with egg volume. It has been shown that changes in environmental conditions (e.g. temperature, salinity) along a latitudinal gradient, can generate clines in reproductive characteristics in both free-living and parasitic species.     

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.