Differential utilization of biochemical components during larval development of the spider crab Maja brachydactyla (Decapoda: Majidae)

Changes in lipid class, fatty acid, fat-soluble vitamins, amino acid and minerals were studied during larval development of Maja brachydactyla Balss, 1922 in order to provide information of its ontogeny and evaluate possible dietary deficiencies and constraints. Four different batches were analyzed from hatching to metamorphosis using enriched Artemia as food. Cultured larvae were in a good nutritional condition as confirmed by the continuous lipid accumulation throughout ontogeny. A regulation of polyunsaturated fatty acids (PUFA) occurred during development in order to maintain adequate basal levels of ω-3 fatty acids (likely through a retro conversion of C₂₀ and C₂₂ into C₁₈ PUFA). Variations in the tocopherol levels indicated its possible role in PUFA protection against oxidation. Essential amino acid balance during development was not correlated with the Artemia feeding regime, but rather reflected inherent variations of the own species ontogeny. Larval requirements in essential minerals were fully supplied by the enriched Artemia.     

Facultative lecithotrophy during larval development of the burrowing shrimp Callianassa tyrrhena (Decapoda: Callianassidae)

Survival, developmental and consumption rate (Artemia nauplii ingested per day) as well as predation efficiency (ingested per available Artemia nauplii) were studied during the larval development of the shallow-water burrowing thalassinid Callianassa tyrrhena (Petagna, 1792), which exhibits an abbreviated type of development with only two zoeal stages and a megalopa. The larvae, hatched from berried females from S. Euboikos Bay (Aegean Sea, Greece), were reared at 10 temperature–food density combinations (19 and 24 °C; 0, 2, 4, 8 and 16 Artemia nauplii d⁻¹). Enhanced starvation resistance was evident: 92 and 58% of starved zoeas I molted to zoea II, while metamorphosis to megalopa was achieved by 76 and 42% of the hatched zoeas at 19 and 24 °C, respectively. The duration of both zoeal stages was affected by temperature, food density and their interaction. Nevertheless, starvation showed different effects at the two temperatures: compared to the fed shrimp, the starved zoeae exhibited accelerated development at 19 °C (8.4 d) but delayed metamorphosis at 24 °C (5.9 d). On the other hand, both zoeal stages were able to consume food at an increased rate as food density and temperature increased. Predation efficiency also increased with temperature, but never exceeded 0.6. Facultative lecithotrophy, more pronounced during the first zoeal stage of C.tyrrhena, can be regarded as an adaptation of a species whose larvae can respond physiologically to the different temperature–food density combinations encountered in the wide geographical range of their natural habitat. Received: 28 February 1998 / Accepted: 21 October 1998     

Changes of respiration and biomass of spider crab (Hyas araneus) larvae during starvation

The influence of starvation on respiration (R), dry weight (W), carbon (C), nitrogen (N), hydrogen (H), and energy content (E; calculated fromC) of spider crab (Hyas araneus L.) larvae was studied in the laboratory. In all larval stages (zoea I and II, megalopa)W increased during postmoult, independent of food, and decreased subsequently. The final reduction inW after continued starvation increased from stage to stage (9, 13, and 20% respectively), but it was always much lower than the decrease inC (44 to 52%),N (42 to 46%),H (50 to 58%), andE (53 to 62%). Individual (R) and weight-specific respiration rates (QO ₂) were reduced by 83 to 88%. The time-dependence of these reductions in metabolism and biomass as well as the rates of change in all parameters studied were described with non-linear regression models and differential equations, respectively. Rates and total amounts of energy lost during starvation were independently calculated fromC andR values, and similar results were mostly obtained. Only in the megalopa stage was there a conspicuous difference between the two estimates: higher losses were calculated fromR. This shows that further (biochemical) data are required for a more complete understanding of the energetics of this stage. Estimates of total protein (fromN) and lipid (fromC) suggest that both constituents serve as metabolic substrates during starvation, but most of the energy originates from the breakdown of protein.Supported by the Deutsche Forschungsgemeinschaft (An 145/1-1)     

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.     

Relationships between sponges and crabs: patterns of epibiosis on Inachus aguiarii (Decapoda: Majidae)

The association between sponges and the crab Inachus aguiarii Brito Capello, 1876 was studied by analysing the relationships between sponge distributional patterns on the crab carapaces and several morphological and biological crab parameters. Juveniles, mature females and mature males were differentiated on the basis of sex dimorphism and terminal pubertary moult. All three groups were fouled to different degrees by sponges. Percent sponge cover was related to carapace size only in mature males whereas mature females, all but one of which were ovigerous, were extensively covered regard-less of their size. It is proposed that some behavioural patterns unique to females, such as long resting periods in sponge-rich microhabitats, are responsible for these high sponge covers in females. Sponges showed two trends in the colonization of the carapace, leading to either a monopolizing or a sharing of the available carapace surface. The sponge species found on the carapaces studied are not obligatory epibionts of crabs, but are believed to reflect the sponge population characterizing the crab home range.     

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.     

Natural diet of larval Penaeus merguiensis (Decapoda: Penaeidae) and its effect on survival

The relationship between Penaeus merguiensis protozoea larvae and their phytoplankton diet was examined using seasonal plankton surveys and in situ rearing experiments. Larval abundance, phytoplankton community structure, and chlorophyll a concentration in Albatross Bay, Gulf of Carpentaria, were monitored monthly for 2 yr. Larval abundance peaked in November (spring) and March (autumn), at which times diatoms were the most abundant group in net samples of phytoplankton and in the guts of larvae. During November 1989 and March 1990, larvae were reared in nylon mesh enclosures positioned throughout the water column at three depths: 0 to 3 m, 3 to 6 m and 6 to 9 m. Overall, larval survival and gut fullness were both higher in November than in March. In both months, larval survival was lower at the surface than at other depths. This correlated with lower chlorophyll a concentrations, but lower total cell densities were not detected. During the in situ experiments, diatoms were the most abundant phytoplankton group in the water column and in the guts of larvae and, therefore, appeared to be the principal diet of larvae. Pigment analysis demonstrated that while gut contents generally reflected the composition of the phytoplankton community, the larvae were not feeding exclusively on diatoms. They also ingested green algae and possibly seagrass detritus. The in situ experiments demonstrated that the predominantly diatom flora in Albatross Bay can provide a nutritionally adequate environment for prawn larvae even at seasonally low levels. It is unlikely, therefore, that starvation is a major cause of mortality of P. merguiensis larvae during either of the biannual peaks in their abundance in Albatross Bay, Gulf of Carpentaria.     

Regulation of respiration during juvenile development of Sepia officinalis (Mollusca: Cephalopoda)

A culture of juvenile Sepia officinalis L. was kept during summer 1985 in the aquaria of the Station Marine, Wimereux, France. During the first four months of juvenile development, oxygen consumption under increasing hypoxia was measured with a closed respirometer. The experiments revealed a high regulatory capacity of juvenile S. officinalis. The critical oxygen concentrations were calculated and their ontogenetical evolution was studied. The critical oxygen concentration increased with increasing development. A linear relationship emerged between the critical oxygen concentration and the logarithm of the wet weight [CO_c (mg O₂ l^-1)=-0.393+0.893×log₁₀(W _w )]. The decreasing regulatory capacity of growing S. officinalis is most probably related to adaptations to a changing ecological environment during development. Another possibility is a physiological change, most probably related to the shift from embryonic to adult hemocyanin.     

Mineral concentration in tissues during ovarian development of the white shrimp Penaeus vannamei (Decapoda: Penaeidae)

Levels of calcium, magnesium, zinc, copper, iron, and manganese were examined in the muscle, hepatopancreas, and gonad of white shrimp, Penaeus vannamei, females at different stages of ovary maturation. Eyestalk-ablated shrimp were sampled during the intermolt stage and classified using the following maturation stages evaluated histologically: cortical stage, vitellogenesis, previtellogenesis, and without gonadic development. In muscle, the copper concentration was significantly lower in the vitellogenic stage than in other stages of ovary development. In the hepatopancreas, manganese levels were higher in the cortical and vitellogenic stages than in animals in previtellogenesis and without gonadic development. No other influence of ovary maturation on muscle and hepatopancreas mineral content was found. In contrast, in the ovary, significant differences of some mineral concentrations were detected. Zinc, iron, and magnesium concentrations were significantly increased in the cortical stage compared to the vitellogenic and previtellogenic stages. Our results indicate that these elements are selectively accumulated during ovary development, especially when the ovaries reach the cortical stage. Received: 14 March 2000 / Accepted: 30 November 2000     

Comparative studies of carotenoids in the Fauna of the Gullmar fjord (Bohuslän,Sweden). II. Crustacea: Eupagurus bernhardus,Hyas coarctatus and Upogebia deltaura

The carotenoids in the crustaceans Eupagurus bernhardus, Hyas coarctatus and Upogebia deltaura from the Gullmar fjord (Bohuslän, Sweden) were investigated. Their presence was determined by means of columnar and thin-layer chromatography. The following carotenoids were found — in E. bernhardus: ß-carotene, canthaxanthin, lutein epoxy, 4-keto-4-ethoxy-ß-carotene, zeaxanthin, astaxanthin and astacene; in H. coarctatus: cryptoxanthin, 4-keto-4-hydroxy-ß-carotene, lutein (pure and epoxy), astaxanthin, and astacene; in U. deltaura: γ-carotene, canthaxanthin, gazaniaxanthin?, isozeaxanthin, cryptoxanthin, 4-keto-4-hydroxy-ß-carotene, zeaxanthin, astaxanthin and astacene.     

An unusual abbreviated larval life in the estuarine burrowing prawn Callianassa kraussi (Crustacea: Decapoda: Thalassinidea)

The larval stages of Callianassa kraussi Stebb. were previously unknown. Larvae were bred out in the laboratory and found to be incapable of swimming. There are two larval stages, lasting 3 to 5 days in sea water of 34 to 35 S at 20 °C. Observations under simulated field conditions in the laboratory, and in the field—where burrow casts were made using a polyester resin, indicated that the larvae metamorphose to the first post-larval stage in the parent burrow, and then burrow directly into the wall of the parent burrow.     

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.     

Elemental composition (C,N,H) in larval and crab-1 stages of Pagurus bernhardus (Decapoda,Paguridae) and Carcinus maenas (Decapoda,Portunidae)

Analyses of individual content of carbon (C), nitrogen (N), and hydrogen (H) were carried out for all larval stages of Pagurus bernhardus and Carcinus maenas, and for newly metamorphosed crabs. Maximum range in total larval development is 12.8 to 165.8 g C, 3.2 to 35.1 g N, and 1.9 to 24.9 g H in P. bernhardus and 3.1 to 43.2 g C, 0.7 to 10.1 g N, and 0.4 to 6.3 g H in C. maenas. From these data energy equivalents were calculated. Maximum range in total larval life is 0.43 to 6.38 J ind. ^-1 in P. bernhardus and 0.1 to 1.49 J ind. ^-1 in C. maenas. There is a 32.4% mean loss of energy in P. bernhardus megalopa development; this seems to describe the normal developmental pattern in this stage. Biomass was determined as fresh and dry weight respectively. Individual dry weight is about 3.6 to 5.6 times higher in P. bernhardus (44 to 340 g) than in C. maenas (12 to 93 g) larvae.Contribution to research project Experimentelle marine Ökosystemanalyse sponsored by Bundesministerium für Forschung und Technologie, Bonn (Grant No. MFU-0328/1)     

Reproduction and larval development of Marenzelleria viridis (Polychaeta: Spionidae)

The pelagic larvae of the polychaete Marenzelleria viridis (Verrill, 1873) are newcomers among the meroplankton of the Baltic Sea, where they sometimes achieve an abundance of up to 21x10⁶ ind m^-3 near the coast, especially in late autumn and even in winter. Benthos samples were collected in the Darss-Zingst bodden chain from April 1992 to 1993 and used to ascertain the reproductive stages of individuals. Development from fertilized egg to benthic juvenile is described on the basis of field material. Details of the morphology of various pelagic stages and the young benthic worm are presented as drawings and scanning electron microscope photographs. In 1992, development of the gametes started in mid-May. The individuals reached maturity in late September after about 20 wk. The first planktonic stage was the fertilized egg. The larvae have initially one and later two pairs of black eyes and, in the 2nd segment, a ciliated pit. Gastrotrochs are present on the third and thereafter every alternate segment. Neuropodial setae develop once the 7-setiger stage is reached. Palps appear at the 10-setiger stage, and neuropodial bidentate hooks from the 10th to 11th setiger on. Metamorphosis into the juvenile benthic stage takes place at the earliest when the 15-setiger stage has been reached. The reproductive seasons of various populations of M. viridis and the developmental patterns of their larvae are discussed, and differences are compared with relevant findings for the spionids.     

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.     

Nucleic acids and growth of larval and early juvenile spider crab,Hyas araneus

The spider crabHyas araneus (L.) was collected from the North Sea in winter 1986–1987 and reared in the laboratory from hatching of the Zoea I (ZI) through the first juvenile instar (CI). Within a given moult cycle, individuals of the same age were sampled in intervals of 2 (ZI, ZII, CI) or 3 d (megalopa) for analysis of dry weight (W), carbon (C), nitrogen (N), hydrogen (H), protein, DNA, and RNA. Lipid was calculated from C. Biomass, growth rate and nucleic acid contents showed high variability during each moult cycle and between instars. Instantaneous growth rates of C were high in postmoult and intermoult, and low in the premoult period of each moult cycle. A shift was observed from high rates of lipid accumulation in the postmoult and intermoult stages to proportionally increasing protein accumulation during late premoult (ZI), or throughout a major part of the remaining moult cycle (in all other instars). DNA was accumulated throughout the ZI and ZII instars, but decreased in late premoult megalopa. It increased again from late intermoult through intermediate premoult in juveniles. RNA increased continuously during ZI and ZII, and decreased in the megalopa, almost to levels that had been found immediately after hatching. In juveniles, variation in RNA followed closely those in DNA. Cell multiplication (expressed by DNA increase) dominated over increase in cell size (defined by the C/DNA ratio) during the zoeal instars and in postmoult through early intermoult in the megalopa and CI. When specific (C-related) RNA values and RNA/DNA ratios were compared with instantaneous growth rates in C and N, no general correspondence was detected. The only significant relationship between specific RNA values and instantaneous C or N growth rates was found in the megalopa. The same held for the relationship between the RNA/DNA ratio and growth. Here, in addition to the megalopa, a correspondence with C growth was also found in the CI instar. Our results suggest that variation in nucleic acids may provide useful insights into mechanisms of growth on the cellular level (cell multiplication vs cell enlargement). However, lack of general correlation with variation in growth rates ofH. araneus larvae shows that the use of nucleic acids as a measure of growth is probably based upon too simplistic assumptions; it may not yield reliable predictions, when growth is associated with developmental events.     

Reproductive behaviour and larval development of Tanais cavolinii (Crustacea: Tanaidacea)

Mature female stages of Tanais cavolinii Milne Edwards are described according to degree of oostegite development. Sexual dimorphism of the chelipeds of copulatory males is explained functionally. An aberrant marsupium consisting of two closed ovisacs with a small slit for sperm transfer protects the brood from sudden unpredictable changes in salinity. Behavioural traits, courtship, copulation, and parental care are described and compared to those of Heterotanais oerstedi Kröyer. The different habitats selected by the two species are reflected by several adaptations in their morphology and behaviour as tube dwellers. A phenomenon unique among invertebrates is that the female T. cavolinii supplies additional yolk to the larvae just before their release. The production and timing of this food supply is described and its adaptive significance is discussed.     

Functional cytology of the hepatopancreas of Penaeus semisulcatus (Crustacea: Decapoda) during the moult cycle

In the shrimp Penaeus semisulcatus, the digestive gland functions as the main site for the digestion, absorption and storage of nutrients and metals during the feeding stages of intermoult and for the mobilization of reserves during the non-feeding stages of pre-and post-moult. These different activities are reflected directly in the cytology of the gland. All the epithelial cells show marked fluctuations in frequency and changes in structure, form and function which occur in phase with the moult cycle. The shrimp were collected offshore from Kuwait in 1980.     

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).