Moult cycle and morphogenesis inHyas araneus larvae (decapoda,majidae), reared in the laboratory

Moult cycle and morphogenesis in larval instars (zoea I, zoea II, megalopa) of the spider crabHyas araneus (L.) were studied in the laboratory. Changes in the epidermis and cuticle were documented photographically at daily intervals to characterize the stages of the moult cycle. Stage A (early postmoult) is a very short period during which the larva takes up water. During late postmoult (B) and intermoult (C) the endocuticle is secreted, and there is conspicuous epidermal tissue condensation and growth. The onset of early premoult (D₀) is characterized by epidermal apolysis, occurring first at the bases of the setae in the telson of zoeal instars or in the rostrum of the megalopa, respectively. Intermediate premoult (D₁) is the main period of morphogenesis, in particular of setogenesis: in the setae of the zoeal telson and carapace there is invagination or (in the zoea II) degeneration of epidermal tissues. Formation of new setae in the interior of epidermal tubules was observed in zoeal maxillipeds and in the antennae of the zoea II and megalopa instars. During late premoult (Stages D_2–4) part of the new cuticle is secreted, and the results of morphogenesis become clearly visible. For technical reasons (rigid exoskeleton) only a preliminary account of the moult cycle in the megalopa can be given. A time schedule is suggested for the stages of the moult cycle. It is estimated that postmoult (A–B) takes ca 9 to 15 % of total instar duration, intermoult (C) ca 22 to 37 %, and premoult (D) ca 48 to 69 %. There is an increasing trend of relative portions of time (% of total instar duration) from instar to instar in Stages A–C (mainly in the latter) and a decreasing trend in Stage D (mainly in D₀ and D_2–4).     

Physiological and biochemical changes during the larval development of a brachyuran crab reared under constant conditions in the laboratory

Larvae of the spider crabHyas araneus were reared in the laboratory at constant conditions (12°C; 32‰S), and their feeding rate (F), oxygen consumption (R), nitrogen excretion (U), and growth were measured in regular intervals of time during development from hatching to metamorphosis. Growth was measured as dry weight (W), carbon (C), nitrogen (N), hydrogen (H) protein, and lipid. All these physiological and biochemical traits revealed significant changes both from instar to instar and during individual larval moult cycles. AverageF was low in the zoea I, reached a maximum in the zoea II, and decreased again in the megalopa. In the zoeal instars, it showed a bell-shaped pattern, with a maximum in the middle (zoea I) or during the first half of the moult cycle (zoea II). MaximumF in the megalopa was observed still earlier, during postmoult. Respiration (R) increased in the zoeal instars as a linear function of time, whereas it showed a sinusoidal pattern in the megalopa. These findings on variation inF andR during larval development confirm results obtained in previous studies onH. araneus and other decapod species. Excretion (U) was measured for the first time with a high temporal resolution in crab larvae. It showed in all three larval instars a bell-shaped variation pattern, with a maximum near the middle of the moult cycle, and significantly increasing average values from instar to instar. The atomicO/N ratio followed an inverse pattern, suggesting a maximum utilization of protein as a metabolic substrate during intermoult. Growth data from the present study and from a number of previous studies were compiled, showing consistency of growth patterns, but a considerable degree of variability between larvae from different hatches reared under identical conditions. The data show the following consistent tendencies: during the first part of each larval moult cycle (in postmoult, partly in intermoult), lipids are accumulated at a higher rate than protein, whereas an inverse growth patterns is typical of the later (premoult) stages. These two different growth phases are interpreted as periods dominated by reserve accumulation in the hepatopancreas, and epidermal growth and reconstruction (morphogenesis), respectively. Differences between individual larval instars in average biochemical composition and growth patterns may be related to different strategies: the zoeal instars and the early megalopa are pelagic feeding stages, accumulating energy reserves (principally lipids) necessary for the completion of larval development, whereas the later (premoult) megalopa is a semibenthic settling stage that converts a significant part of this energy to epidermal protein. The megalopa shifts in behaviour and energy partitioning from intense feeding activity and body growth to habitat selection and morphogenesis, preparing itself for metamorphosis, i.e. it shows an increasing degree of lecithotrophy. Data from numerous parallel elemental and biochemical analyses are compiled to show quantitative relationships betweenW, C, N, H, lipid, and protein. These regressions may be used as empirical conversion equations for estimates of single chemical components in larvalHyas araneus, and, possibly, other decapods.     

Growth patterns,chemical composition and oxygen consumption in early juvenileHyas araneus (Decapoda: Majidae) reared in the laboratory

Early (instar I and II) juveniles of the spider crabHyas araneus were reared under constant conditions (12 °C, 32‰S) in the laboratory, and their growth, biochemical composition, and respiration were studied. Every second day, dry weight (W), ash-free dry weight (AFW), and contents of ash, organic and inorganic carbon (C), nitrogen (N), hydrogen (H), protein, chitin, lipid, and carbohydrates were measured, as well as oxygen consumption. Changes in the absolute amounts of W. AFW, and C, N, and H during the moulting cycle are described with various regression equations as functions of age within a given instar. These patterns of growth differ in part from those that have been observed during previous studies in larval stages of the same and some other decapod species, possibly indicating different growth strategies in larvae and juveniles. There were clear periodic changes in ash (% of W) and inorganic C (as % of total C), with initially very low and then steeply increasing values in postmoult, a maximum in intermoult, and decreasing figures during the premoult phase of each moulting cycle. Similar patterns were observed in the chitin fraction, reaching a maximum of 16% of W (31% of AFW). Ash, inorganic C, and chitin represent the major components of the exoskeleton and hence, changes in their amounts are associated with the formation and loss of cuticle material. Consequently, a high percentage of mineral matter was lost with the exuvia (76% of the late premoult [LPM] ash content, 74% of inorganic C), but relatively small fractions of LPM organic matter (15% of AFW, 11% of organic C, 5–6% of N and H). These cyclic changes in the cuticle caused an inverse pattern of variation in the percentage values (% of W) of AFW, organic C, N, H, and biochemical constituents other than chitin. When these measures of living biomass were related to, exclusively, the organic body fraction (AFM), much less variation was found during individual moulting cycles, with values of about 43–52% in organic C, 9–10% in N, 6–9% H, 31–49% of AFW in protein, 3–10% in lipid, and <1% in carbohydrates. All these constituents showed, on the average, a decreasing tendency during the first two crab instars, whereas N remained fairly constant. It cannot be explained at present, what other elements and biochemical compounds, respectively, might replace these decreasing components of AFW. Decreasing tendencies during juvenile growth were observed also in the organic C/N and in the lipid/protein weight ratios, both indicating that the proportion of lipid decreased at a higher rate than that of protein. Changes were observed also in the composition of inorganic matter, with significantly lower inorganic C in early postmoult (2–4% of ash) than in later stages of the moult cycle (about 9%). This reflected probably an increase in the degree of calcification, i.e. in the calcium carbonate content of the exoskeleton. As a fraction of total C, inorganic C reached maximum values of 17 and 20% in the crab I and II instars, respectively. The energy content of juvenile spider crabs was estimated independently from organic C and biochemical constituents, with a significant correlation between these values. However, the former estimates of energy were, on the average, significantly lower than the latter (slope of the regression ≠1). Since organic C should be a reliable integrator of organic substances, but the sum of protein, lipid, chitin, and carbohydrates amounted to only 60–91% of AFW, it is concluded that the observed discrepancy between these two estimates of energy was caused by energy from biochemical constituents that had not been determined in our analyses. Thus, energy values obtained from these biochemical fractions alone may underestimate the actual amount of organic matter and energy. Respiration per individual in juvenile spider crabs was higher than that in larval stages of the same species (previous studies), but their W-specific values of oxygen consumption (QO₂) were lower than in conspecific larvae (0.6–2μg O₂·[mg W]⁻¹). QO₂ showed a consistent periodic pattern in relation to the moult cycle: maximum values in early postmoult, followed by a rapid decrease, and constant values in the intermoult and premoult phases. This variation is interpreted as an effect mainly of cyclic changes in the amounts of cuticle materials which are metabolically inactive. From growth and respiration values (both expressed in units of organic C), net growth efficiency, K₂, values may be calculated. In contrast to previous findings in larval stages, K₂ showed an increasing trend during growth of the first two juvenile instars ofH. araneus.     

Growth and reproduction of eyestalk ablated penaeus canaliculatus (Olivier, 1811) (crustacea:Penaeidae)

The effects of single (unilateral) eyestalk ablation on the growth and reproduction of male and female Penaeus canaliculatus (Olivier) were compared with those of unablated (control) individuals. Prawns ≤ 10 mm in carapace length ablated in the premoult stage suffered high mortality. Prawns recognized as immature when ablated always moulted irrespective of their moulting stage; ovaries in females did not become vitellogenic nor did spermatogenesis occur in males. Mature females ablated in the premoult stage underwent moulting while those in the postmoult stage developed mature ovaries. Mature males in the postmoult or intermoult stages took longer to moult than those that were in the premoult stage when ablated. The Von Bertalannfy equations describing growth in P. canaliculatus were as follows: L_t = 25.6 [1−e^{−0.0756(t−t_o)}]for ablated males; L_t = 25.3 [1−e^{−0.059(t−t_o)}] for unablated males; L_t= 37.2 [1−e^{−0.048(t−t_o)}] forablated females; L_t = 33.4 [1−e^{−0.044(t−t_o)]} for unablated females. Differences in the growth rates were a result of both the moulting frequency and the increment in size at moult. However, the relative contribution of these two factors to growth varied with sex as well as with size. In both sexes, ablated individuals became sexually mature earlier; females spawned earlier. Although moulting frequency and the total number of spawns were greater for ablated females, the mean number of eggs produced (per spawn as well as total) by unablated females was higher and the mean hatching success was better.     

Modelling developmental changes in the carbon and nitrogen budgets of larval brachyuran crabs

The uptake and partitioning of nutritional carbon (C) and nitrogen (N) were studied during the complete larval development of a brachyuran crab,Hyas araneus, reared under constant conditions in the laboratory. Biochemical and physiological data were published in a foregoing paper, and complete budgets of C and N were now constructed from these data. Regression equations describing rates of feeding (F), growth (G), respiration (R), and ammonia excretion (U) as functions of time during individual larval moult cycles were inserted in a simulation model, in order to analyse time-dependent (i.e. developmental) patterns of variation in these parameters as well as in bioenergetic efficiencies. Absolute daily feeding rates (F; per individual) as well as carbon and nitrogen-specific rates (F/C, F/N) are in general maximum in early, and minimum in late stages of individual larval moult cycles (postmoult and premoult, respectively). Early crab zoeae may ingest equivalents of up to ca 40% body C and 30% body N per day, respectively, whereas megalopa larvae usually eat less than 10%. Also growth rates (G; G/C, G/N) reveal decreasing tendencies both during individual moult cycles and, on the average, in subsequent instars. Conversion of C and N data to lipid and protein, respectively, suggests that in all larval instars there is initially an increase in the lipid: protein ratio. Protein, however, remains clearly the predominant biochemical constituent in larval biomass. The absolute and specific values of respiration (R; R/C) and excretion (U; U/N) vary only little during the course of individual moult cycles. Thus, their significance in relation toG increases within the C and N budgets, and net growth efficiency (K ₂) decreases concurrently. Also gross growth and assimilation efficiency (K ₂; A/F) are, in general, maximum in early stages of the moult cycle (postmoult). Biochemical data suggest that lipid utilization efficiency is particularly high in early moult cycle stages, whereas protein utilization efficiency is higher in later stages. Only the zoea II appears to accumulate lipid from food constantly with a higher conversion efficiency than protein. The cumulative C and N budgets show in subsequent larval instars conspicuously increasing figures in all of their parameters.F andG increase to a particularly high extent from the first to the second zoeal instar, whereasR, U, exuvia production (G _E), and total assimilation (A) reveal a greater increase from the zoea II to the megalopa. Respiratory, excretory, and exuvial losses increase in subsequent larval instars at higher rates than tissue growth and, hence,K ₂ decreases in the same order. In the C budget,K ₂ values of 0.63 (zoea I). 0.56 (zoea II), and 0.29 (megalopa) were calculated (or: 0.56, 0.46, and 0.16 after subtraction of exuviae). In the N budget, corresponding values of 0.76, 0.66, and 0.45 (or: 0.72, 0.62, 0.38 without exuviae) were obtained. AlsoK ₁ decreases slightly in subsequent instars, whereasA/F reveals rather an increasing tendency, at least from the zoeal instars to the megalopa. Changes in the uptake and partitioning of matter in crab larvae are discussed in relation to developmental events and changes in life style before metamorphosis.     

Stimulation of metamorphosis in an estuarine crab, Chasmagnathus granulata (Dana, 1851): temporal window of cue receptivity

The larvae of most benthic marine invertebrate species must develop for a minimum of time in the plankton before they become competent for settlement and metamorphosis in response to stimulating external cues. In an experimental laboratory study, we identified the temporal window of cue receptivity within the moulting cycle of the megalopa stage of an estuarine crab, Chasmagnathus granulata Dana. This species shows an export strategy including an early larval transport to coastal marine waters where zoeal development takes place, followed by the return of the megalopa stage to brackish habitats where the adults live. In two series of experiments (A, B), megalopae were exposed for differential periods to a combination of metamorphosis-stimulating cues which had previously been found effective (seawater conditioned with adult odor and presence of mud). In experimental series A, these cues were added on successively later days of the moulting cycle, while series B comprised treatments in which the cues were provided from the first day (postmoult) and removed on successively later days of the moulting cycle. Each series of experiments was repeated with larvae originating from two different females (F1, F2). The average development time of megalopae kept continuously in the presence of cues (control experiments, C1) ranged in the two hatches from 9.3 to 9.6 days. In the inverse controls where no cue was added at any time (C2), megalopal development to metamorphosis took on average 11.2-12.0 days. In series A, development duration in treatments with exposure to the cues commencing within 3-4 days after moulting was not significantly different from that in the permanently exposed C1 controls. Later beginning of the exposure, by contrast, had no stimulating effect (significant delay compared to C1, no significant difference from unexposed control, C2). In series B, no significant differences in development time were observed between the C1 controls and treatments with an initial exposure for a minimum of 4 or 6 days of the moulting cycle (F1, F2, respectively). Shorter initial periods of exposure had no metamorphosis-stimulating effects (no significant difference from C2). In conclusion, our results from both experiments suggest that the megalopa stage of C. granulata is most receptive of stimulating cues during a period lasting from ca. one third to one half of the moulting cycle, which coincides with the transition between stages C (intermoult) and D₀ (early premoult) of Drach's classification system. This suggests an interaction of extrinsic stimulating cues with intrinsic (hormonal) factors involved in the control of the moulting cycle.     

Effects of reduced salinities on development and bioenergetics of early larval shore crab, Carcinus maenas

The shore crab, Carcinus maenas L. (Portunidae), is a coastal and estuarine species, which can live and reproduce under brackish water conditions; freshly hatched larvae have been observed in the field at salinities below 15‰. In the present laboratory study, the tolerance of hypo-osmotic stress was experimentally investigated in early larvae of a marine (North Sea) population of C. maenas reared at four different salinities (15, 20, 25, 32‰). Two and 4 days after hatching, the Zoea I larvae were moult-staged microscopically, and their rates of respiration and growth (changes in dry weight, W, carbon, C, nitrogen, N, and hydrogen, H) were measured. Survival and development were monitored until the megalopa was reached: 15‰ did not allow for development beyond the first zoeal stage, while metamorphosis to the megalopa was reached at salinities ≥20‰. At 20‰, development was significantly delayed and mortality enhanced as compared with 25 and 32‰. Rates of growth and respiration decreased during exposure to reduced salinities ≤25‰. Hence, the suppression of growth could not be explained as a consequence of enhanced metabolic losses per larva. Instead, a partial C budget indicates that the Zoea I larvae suffered from decreased capabilities of assimilating ingested and subsequently converting assimilated matter to tissue growth. Net growth efficiency (K₂, C-based) was at 25 and 32‰ initially high (>60% during the postmoult and intermoult stages of the Zoea I moult cycle), but decreased during the later stages (down to ≤30% in premoult). An inverse pattern of C partitioning was observed at ≤20‰, with initially low K₂ values (≤21% during the first 2 days of the moult cycle), and a later increase (up to ≥46% in premoult). Thus, larval growth was initially suppressed under conditions of reduced salinity, but this was later (during premoult) partially compensated for by an increase in C assimilation and K₂. Our observations indicate that Zoea I shore crab larvae react during the late stages of their moulting cycle less sensitively against reduced salinities than during postmoult and intermoult. This suggests that the transition between moult cycle stages C and D₀ may be a critical point for effects of hypo-osmotic stress, similarly as already known in relation to effects of nutritional stress. Negative effects were found also when freshly hatched Zoea I shore crab larvae were exposed only transitorily (for 24–72 h) to 20‰, with significantly lower rates of survival, development, growth, respiration, and K₂. These effects increased with increasing duration of initial exposure to reduced salinity.     

Bioenergetics of abbreviated larval development in the bromeliad crab,Metopaulias depressus (Decapoda: Grapsidae)

• 1.1. Larvae of the bromeliad crab, Metopaulias depressus Rathbun, were reared in the laboratory, and changes in dry weight (W), ash-free W (AFW), carbon, nitrogen, hydrogen, protein, lipid, carbohydrates and respiration rate were measured during development from hatching to metamorphosis.
• 2.2. Development was successful in rain-water from bromeliads (pH < 5–6), but not in river water from the same region (pH 8). It is abbreviated, with two non-feeding zoeal stages (2.5–3.5 days each) and a feeding megalopa (8.5–10 days). Development to metamorphosis can also be completed in the absence of food (facultative lecithotrophy).
• 3.3. Dry weight and other absolute biomass values per individual vary significantly between different hatches, whereas changes in the relative (% of W or AFW) composition follow quite invariable patterns: ash increases from hatching through the first part of megalopa development, organic biomass decreases concurrently.
• 4.4. Elemental and biochemical data show that lecithotrophy of the zoeal stages as well as continued endotrophic development in the megalopa depend chiefly on degradation of lipid reserves and less on protein. No significant growth was observed in organic constituents when food was available, but without food the megalopa reached metamorphosis with only half the lipid and less than two thirds the protein of fed siblings.
• 5.5. The relationship between C and lipid is similar in M. depressus larvae as in planktotrophic marine crab larvae, whereas that between N and protein differs; it indicates the presence of unusually large quantities of unidentified non-protein N.
• 6.6. Exuvial losses of late premoult biomass or energy are very low in the zoeal stages (2 and 3%), but increase in the megalopa (16% in W, 10% in C, 7–8% in N, H and energy).
• 7.7. Respiration rate per individual increases gradually during larval development (0.6–0.8 μg O₂/hr). Starved megalopa larvae reveal lower individual but higher W-specific metabolism than fed larvae.
• 8.8. Bioenergetic traits of abbreviated larval development are discussed in relation to those known from regular (planktotrophic marine) development of brachyuran crabs. M. depressus is highly adapted to life and development in a physically extreme terrestrial environment.

     

Partitioning and utilization of energy during the larval development of the xanthid crab, Rhithropanopeus harrisii (Gould)

An energy budget is constructed for the larval development of the crab Rhithropanopeus harrisii (Gould) fed nauplii of the brine shrimp Artemia salina (L.). Between the first zoeal instar and the megalopa, there is a 5.4-fold increase in caloric consumption and a 13.2-fold increase in dry weight. Weight specific energy content increases through the zoeal stages and drops at the megalopa. Rate of oxygen consumption increases steadily throughout development. Assimilation, gross growth, and net growth efficiencies increase steadily through zoeal development and drop at the megalopa. Calories put into tissue production exceed those expended via respiration in zoeal stages II–IV; the reverse is true in zoeal stage I and the megalopa.

A total energy budget has been calculated and the partitioning of energy is discussed in relation to other physiological studies on this species.     

Influence of temperature on larval survival, development, and respiration inChasmagnathus granulata (Crustacea, Decapoda)

Larvae of an estuarine grapsid crabChasmagnathus granulata Dana 1851, from temperate and subtropical regions of South America, were reared in seawater (32/%.) at five different constant temperatures (12, 15, 18, 21, 24 °C). Complete larval development from hatching (Zoea I) to metamorphosis (Crab I) occurred in a range from 15 to 24 °C. Highest survival (60% to the first juvenile stage) was observed at 18 °C, while all larvae reared at 12 °C died before metamorphosis. The duration of development (D) decreased with increasing temperature (T). This relationship is described for all larval stages as a power function (linear regressions after logarithmic transformation of bothD andT). The temperature-dependence of the instantaneous developmental rate (D ⁻¹) is compared among larval stages and temperatures using the Q₁₀ coefficient (van't Hoff's equation). Through all four zoeal stages, this index tends to increase during development and to decrease with increasingT (comparing ranges 12–18, 15–21, 18–24 °C). In the Megalopa, low Q₁₀ values were found in the range from 15 to 24 °C. In another series of experiments, larvae were reared at constant 18 °C, and their dry weight (W) and respiratory response to changes inT were measured in all successive stages during the intermoult period (stage C) of the moulting cycle. Both individual and weight-specific respiration (R, QO ₂) increased exponentially with increasingT. At each temperature,R increased significantly during growth and development through successive larval stages. No significantly differentQO ₂ values were found in the first three zoeal stages, while a significant decrease with increasingW occurred in the Zoea IV and Megalopa. As in the temperature-dependence ofD, the respiratory response to changes in temperature (Q₁₀) depends on both the temperature range and the developmental stage, however, with different patterns. In the zoeal stages, the respiratory Q₁₀ was minimum (1.7–2.2) at low temperatures (12–18 °C), but maximum (2.2–3.0) at 18–24 °C. The Megalopa, in contrast, showed a stronger metabolic response in the lower than in the upper temperature range (Q₁₀=2.8 and 1.7, respectively). We interpret this pattern as an adaptation to a sequence of temperature conditions that should typically be encountered byC. granulata larvae during their ontogenetic migrations: hatching in and subsequent export from shallow estuarine lagoons, zoeal development in coastal marine waters, which are on average cooler, return in the Megalopa stage to warm lagoons. We thus propose that high metabolic sensitivity to changes in temperature may serve as a signal stimulating larval migration, so that the zoeae should tend to leave warm estuaries and lagoons, whereas the Megalopa should avoid remaining in the cooler marine waters and initiate its migration to wards shallow coastal lagoons.     

Tissue accumulation and the moult cycle in juveniles of the Australian freshwater crayfish Cherax destructor

1. This paper investigates moult stage and size-specific changes in whole body composition during growth in juvenile crayfish in order to better describe the nature of growth and energy use. 2. Composition is described in terms of moult stage and size-specific wet, dry and ash-free dry weight, water, carbon, protein nitrogen, non-protein nitrogen, ash and energy. Dry weight and ash-free dry weight (AFDW) peaked in the middle and in the later stages of premoult. Both peaks were about 2–2.3 times postmoult weight. Losses in tissue weight during ecdysis were substantial in the smallest crayfish but declined with size. Water was taken up between late premoult and early postmoult. Tissue accumulation occurred primarily between B and D₁.₃, with further weight gain largely the result of fluid uptake. Ash increased immediately postmoult with a major peak occurring during intermoult. Mean organic carbon varied between 33 and 35.5% of the body and 49% of the exuvia. Chitin varied between 9 and 17% of body AFDW and made up about 50.5% of the exuvia. Protein content varied between 47 and 62% of body AFDW and about 25% of the exuvia. Carbon, chitin and protein were not affected by moult stage but protein declined with ocular carapace length (OCL) in larger crayfish. Mean caloric content varied between 19 and 22 J mg-¹ AFDW depending on size and moult stage. Caloric content increased with OCL during premoult and early intermoult then declined with size until part of the way through premoult. 3. Relationships between protein, chitin and remaining carbon (organic carbon minus chitin) were examined. It is suggested that protein and some carbon are catabolized during the moulting process, possibly to fuel metabolism. Models are presented showing changes in proximate composition over the moult cycle for two sizes of crayfish, and tissue and energy accumulation and loss over a series of moult cycles and sizes from 3.1 mm to 17 mm OCL.     

Larval development of the subantarctic squat lobster Munida subrugosa (White, 1847) (Anomura: Galatheidae), reared in the laboratory

The larval development of the squat lobster Munida subrugosa from subantarctic waters of the Beagle Channel (Tierra del Fuego, Argentina) was studied under controlled laboratory conditions of temperature, salinity, and food supply. Developmental times, survival, and growth of larvae and early juveniles were investigated. Hatching of the entire brood always occurred during one night. Larvae were kept in 100ml individual bowls with filtered seawater at 8 ± 0.5°C and fed with Artemia spp. nauplii three times a week. Larvae passed through 6 zoeal instars and one megalopa. Previously, only five zoeal instars were known from this species. Mean cumulative durations of the zoeal stages I to VI were: 20.5 ± 2.5, 33.9 ± 4.1, 43.3 ± 5.4, 52.6 ± 5, 61.2 ± 3.9, and 83days, respectively. By adding the 28days that a single megalopa took to metamorphose to crab I stage, the complete larval development lasted 111days. Highest mortality occurred prior to the moult from the zoea I to zoea II stage (79.21% ± 18.65%) and during the moult from zoea VI to megalopa (92.86%). Carapace length was 1.64 ± 0.06, 1.52 ± 0.16, 1.57 ± 0.26, 1.64 ± 0.21, 2.11 ± 0.35, and 2.58 ± 0.19mm, for zoeal stages I to VI, respectively. Carapace length of megalopae and crab I instars was similar (2.85 ± 0.28 and the 2.84 ± 0.05, respectively). Unlike other subantarctic decapods, which show a tendency towards abbreviated larval development and/or some degree of endotrophy, M. subrugosa shows an extended planktotrophic larval development synchronized with short seasonal plankton production in austral summers.     

Effect of bilateral eyestalk ablation on ovarian development and moulting in early and late intermoult stages of female spiny lobster Panulirus homarus (Linnaeus, 1758)

Abstract

The present study describes the effect of bilateral eyestalk ablation (BESA) on reproduction and moulting of spiny lobster Panulirus homarus females in their early and late intermoult stages. The lobsters obtained from the wild were conditioned for the experiment. The experiments were conducted at the Calicut research centre of Central Marine Fisheries Research Institute, India. Eyestalk ablation of females was done by ligation. The responses of the bilaterally eyestalk-ablated lobsters were statistically analysed. BESA conducted on the early and late intermoult stages resulted in the simultaneous acceleration of the somatic growth and reproductive processes with higher emphasis for oogenesis in lobsters ablated in the early intermoult phase and comparatively lower activity in those ablated in the late intermoult phase. Ablation in late intermoult phase resulted in faster entry into the premoult stage compared to the control.     

The accumulation of age pigments during larval development of the spider crab,Hyas araneus (Decapoda,Majidae)

• 1.1. Lipofuscin, body carbon and respiration rates were measured in Hyas araneus from hatching to metamorphosis. Lipofuscin was measured spectrofluorometrically from the chloroform phase of chloroform/methanol extracts.
• 2.2. Excitation/emission spectra of both the chloroform and the methanol/aqueous phase showed one distinct fluorescence peak in the chloroform (410–415 nm emission/340–350 nm excitation) and the methanol/aqueous phase (405/350 nm) of zoea I (directly after hatching) and megalopa (0 and 24 days old).
• 3.3. Individual lipofuscin concentrations increased continuously during zoea I and halfway through zoea II, but remained constant through the entire megalopa despite high metabolic activity in this stage.
• 4.4. Individual lipofuscin concentrations were positively correlated with body carbon and carbonspecific lipofuscin was negatively correlated.
• 5.5. Moulting caused considerable loss of lipofuscin. During the first two larval ecdyses 17–18% were lost, with the shed moults containing only 3.4–4.5% of the lipofuscin found in late premoult individuals.
• 6.6. The different patterns of lipofuscin accumulation in respective larval stages is discussed in regard to mitotic activity of tissues. While in the zoea, growth is more related to lipid formation and biomass accumulation, in the megalopa morphogenetic processes require substantial epidermal growth, i.e. protein accumulation. However, the question why in the megalopa no increase in lipofuscin is found, remains unanswered.

     

Moult cycle and its chronology in Mysis mixta and Neomysis integer (Crustacea, Mysidacea): implications for growth assessment

The moult-staging technique was used to determine the main moult stages (MS) and their duration in laboratory-reared juveniles and subadults of Mysis mixta and Neomysis integer. The relative duration of each stage was similar for both species with premoult occupying the major part of the moult cycle (51% and 44% for M. mixta and N. integer, respectively), followed by postmoult (26% and 34%) and intermoult (23% and 22%). Effects of temperature and feeding regimes on the chronology of the moult cycle were investigated. When the duration of the moult cycle (MCD) was extended by manipulating the feeding regime, animals prolonged their late postmoult and early premoult stages. At 12 and 5 °C, no specific moult stage varied in relative duration as long as food supply was high. Field application of moult staging for growth assessment was tested using wild-caught M. mixta. The MCD estimated via moulting experiments was compared to that obtained by analyzing moult stage distribution combined with the experimentally obtained data on stage duration. Close correspondence between two methods of MCD assessment was observed with moult-staging technique being particularly useful in situations when conducting of experiments immediately after collection is not feasible.     

The anal glands of Rhapidostreptus virgator (Diplopoda: Spirostreptidae)

Summary In Rhapidostreptus virgator exocrine gland complexes are found in the anal valves of both sexes. Every gland complex consists of about 200 secretory units, each of which is comprised of four cells: two secretory cells, an intermediary cell, and a canal cell. The amount of secretion produced by these glands varies during the intermoult cycle: it is very small in freshly moulted individuals (postmoult phase), at a medial level during the following intermoult phase, and very large in the premoult phase. The secretion may be used to form the excrement clumps and above all to build the moulting chamber.     

Nutritional requirements during larval development of the portunid crab,Callinectes sapidus Rathbun

Three kinds of diet were used to identify the critical periods of nutritional vulnerability during larval development of the portunid crab, Callinectes sapidus Rathbun. A diet consisting solely of the rotifer Brachionus plicatilis Müller is not sufficient for complete developmetn. Development to metamorphosis can occur if rotifers are replaced by Day 15 with Artemia salina L. nauplii, but a diet of A. salina between Days 15 and 21 is also not sufficient for complete development. Delay in giving a brine shrimp diet beyond Day 15 causes a reduction in survival to the megalopa with an apparent threshold between Days 22 and 29, delay in molting in late instars, and increased frequency of supernumeracy larvae, many of which molt subsequently to the megalopa. Development to the megalopa on the rotifer diet is possible if crab larvae are initially fed a favorable control diet for as little as 14 days after hatching. Extension of time on the control increases survival to the megalopa slightly, has little effect on molt frequency, but reduces the number of zoeal instars.These data are interpreted in the context of identification of the unknown dietary requirement and for its implication to evolution of reproduction in the Brachyura.     

Changes in biomass and chemical composition during lecithotrophic larval development of the southern king crab, Lithodes santolla (Molina)

Changes in biomass and elemental composition (dry mass, W; carbon, C; nitrogen, N; hydrogen, H) were studied in the laboratory during complete larval and early juvenile development of the southern king crab, Lithodes santolla (Molina), formerly known as Lithodes antarcticus (Jacquinot). At 6±0.5 °C, total larval development from hatching to metamorphosis lasted about 10 weeks, comprising three demersal zoeal stages and a benthic megalopa, with mean stage durations of 4, 7, 11 and 47 days, respectively. No differences in development duration or mortality were observed in larvae either fed with Artemia sp. nauplii or unfed, indicating that all larval stages of L. santolla are lecithotrophic. First feeding and growth were consistently observed immediately after metamorphosis to the first juvenile crab stage. Regardless of the presence or absence of food, W, C, N and H decreased throughout larval development. Also the C:N mass ratio decreased significantly, from 7.7 at hatching to 4.1 at metamorphosis, indicating that a large initial lipid store remaining from the egg yolk was gradually utilized as an internal energy source, while proteins played a minor role as a metabolic substrate. In total, 56-58% of the initial quantities of C and H present at hatching, and 20% of N were lost during nonfeeding larval development to metamorphosis. Nine to ten percent of the initially present C, N and H were lost with larval exuviae, half of these losses occurring in the three zoeal stages combined and another half in the megalopa stage alone. Metabolic biomass degradation accounted for losses of about 47-50% in C and H but for only 10% in N. Hence, most of the losses in C and H reflected metabolic energy consumption (primarily lipid degradation), while about half of the losses in N and two thirds of those in W were due to larval exuviation. Complete independence from food throughout larval development is based on an enhanced maternal energy investment per offspring and on energy-saving mechanisms such as low larval locomotory activity and low exuvial losses. These traits are interpreted as bioenergetic adaptations to food-limited conditions in Subantarctic regions, where a pronounced seasonality of day length limits the period of primary production, while low temperatures enforce a long duration of pelagic development.     

Ammonia excretion of the sand-shrimpCrangon crangon (L.) during the moult cycle

Summary Ammonia excretion in the shrimpCrangon crangon (L.) shows a cyclical pattern during the moult cycle. At the early premoult (stage D₀) there is a 7 to 17% increase over that characteristic at intermoult; at late premoult (stage D₂) excretion decreases to a minimum; immediately after ecdysis, the excretion rate is 1.5 to 2.6 times higher than at the intermoult stage (Table 1).These variations appear to be correlated with protein breakdown and protein synthesis, and possibly with periods of high metabolic activity (growth resulting from cell multiplication).Ammonia excretion of animals in standing water is higher than of those in running sea-water. Excretion might possibly be influenced by a semi-lunar rhythm.