Calorific,ash, carbon and nitrogen content in relation to length and dry weight of Parathemisto gaudichaudi (Amphipoda: Hyperiidea) in the North East Atlantic Ocean

The calorific, ash, carbon and nitrogen content, length and dry weight were determined for the hyperiid Parathemisto gaudichaudi (Guerin). Regression equations for all these variables were determined so that they can be estimated by calculation from measurements of length of the hyperiid. Mean values for total nitrogen and carbon were 7.79±0.85% and 36.80±4.18% of the dry weight, respectively. The carbon to calorific equivalent for P. gaudichaudi was 10.37 kcal g^-1 carbon (9.13 kcal g^-1 when corrected for nitrogen). The calorific value for ash-free adult P. gaudichaudi was 5.138 kcal g^-1±1.309 (4.510 kcal g^-1 when corrected for nitrogen). This large variation in the calorific content (coefficient of variation of 25.84%) can be accounted for largely by variation in the ash content (coefficient of variation of 21.84%). The calorific value determined for P. gaudichaudi is similar to that measured for other carnivorous crustaceans and adds support to the hypothesis that animals with high calorific content have a low fecundity and an energy-rich store which can be used as a buffer during unfavourable periods in their life.     

Brood size and chemical composition of Pareuchaeta norvegica (Crustacea: Copepoda) in Loch Etive,Scotland

Brood size and chemical composition (carbon, nitrogen, and ash) of Pareuchaeta norvegica in Loch Etive, Scotland have been determined. From 23 to 37 eggs were found in each egg mass (mean=30 eggs). The eggs are oval; the mean size of the longer axis is 438 m. Variation in egg mass (as dry weight) is related to increase in dry weight of the maternal body. an increase of 1 mg in maternal body dry weight produces an increase of 0.03 mg dry weight in the egg mass. Carbon content increases with increasing body dry weight both in pre-spawning and spent females, but nitrogen content decreases with increasing body weight. Thus, the carbon:nitrogen ratio increases with increasing body dry weight. The carbon content of an egg mass ranges from 61.4 to 65.3%, and nitrogen content from 9.6 to 13.2% of the total dry weight. Some examples of partitioning of chemical components of the maternal body into the egg mass are also described.     

Ecological investigations of the zooplankton community of balsfjorden,northern norway: The genital system in Calanus finmarchicus and the role of gonad development in overwintering strategy

The structure of the reproductive system in Copepodite Stages IV and V of the marine copepod Calanus finmarchicus (Gunnerus) is described, together with seasonal variations in gonad development and sex ratio, from Balsfjorden (69°21N; 19°06E), a subarctic fjord in northern Norway. The genital system in Copepodite Stage IV consists of a small gonad with two genital ducts. The gonad of Copepodite Stage V in its most immature condition resembles that seen in Copepodite Stage IV, whereas the sex of Stage V copepodites with mature gonads can be easily and reliably determined. The genital ducts are the key characters for separating males and females in Copepodite Stage V; potential males have one and potential females two genital ducts connecting the gonad to the genital opening on the first urosome segment. Adults males were rare compared to females, and were only in the majority at the end of January and beginning of February. The sex-ratio also favoured females in Copepodite Stage V. Seasonal peaks in the proportion of adult males reflected peaks of males in Copepodite Stage V, suggesting that seasonal variations in the adult sex-ratio are a direct result of a situation which has already been determined in Copepodite Stage V. The seasonal variation in gonad development in Copepodite Stages IV and V of C. finmarchicus reveals that the size of maturity of the gonad varies throughout the year. This study indicates that visible sex-differentiation in Stage V and moulting into adults occurs in Balsfjorden at least 2 months before spawning in April. Phytoplankton levels are immeasurable before the spring diatom increase starts at the end of March, implying that development and maturation of gonads are dependent upon internal energy resources, whereas the final act of spawning seems to require energy input from phytoplankton.     

Early development and estimated life cycle of the mesopelagic copepod Pareuchaeta elongata in the southern Japan Sea

Life cycle of the mesopelagic copepod Pareuchaeta elongata was investigated combining laboratory rearing data on the eggs, nauplii, and early copepodites with field growth data on the late copepodites. Examination and incubation at near the in situ temperature (0.5°C) of egg sacs collected from the field indicated that the clutch size was 13 to 24 eggs (mean: 20), hatching time was 39.4d (mean), and hatchability was low (mean: 28.5%). The development time at 0.5°C was 2.4 d for Nauplius Stage 1 (N1), 4.6 d for N2, 6.2 d for N3. 7.4 d for N4, 7.1 d for N5, 20.8 d for N6, 36.7 d for Copepodite Stage 1 (C1) and 65.3 d for C2. From the numerical analysis of seasonal samples collected from the field, the development time was estimated as 1 mo for C3, 2 mo for C4 and 2.5 mo for C5. Thus, the egg hatching time plus the integrated development time by stage was 355 d or nearly 1 yr (i.e. span of one life cycle). Duration of the C6 (adult) was estimated as>2.5 mo. Combining the present data on development times of each stage with published data on the major spawning season (August to October) and ontogenetic migration, a schematic representation of the life cycle of this copepod was developed. Between-stage comparison of wet, dry, and ash-free dry weights of all developmental stages of preserved wild specimens revealed that there is almost no gain in weight during naupliar stages, and the greatest weight increment over the life cycle was during the C4 stage. The present results are compared with those for the same and related species living in other regions.     

Dry weight, carbon and nitrogen components of caprellid amphipods (Crustacea) inhabiting the Sargassumyezoense community of Otsuchi Bay, northeastern Japan

Dry weight (DW), carbon (C) and nitrogen (N) contents in four species of caprellids, Caprella danilevskii, C. subinermis, C. penantis R-type, and C. verrucosa, inhabiting Sargassumyezoense (Yamada) in Otsuchi Bay, northeastern Japan, were measured based on live specimens. In all species males tend to grow larger than females in pereonite II length, which is used as an indicator of body length, and have greater dry weight, carbon and nitrogen contents. All three parameters (DW, C and N) had highly significant relationships against pereonite II length. Regressions of female pereonite II length against the three parameters significantly differed from males in the four species. Average percentages of carbon and nitrogen contents of the total dry weight ranged from 30.82 to 32.21% and 5.82 to 6.86%, respectively. The carbon/nitrogen ratio tended to decrease against increasing pereonite II length, e.g., 7.2 to 4.3 in C. danilevskii. Received: 25 November 1996 / Accepted: 31 July 1997     

Ecobiochemical studies on the deep-water pelagic community of Korsfjorden,Western Norway. Adenine nucleotides in zooplankton

Adenine nucleotides of zooplankton were extracted from freeze-dried individuals killed in liquid nitrogen. 96% of the ATP in frozen samples stored at-26°C degraded to AMP in 8 days, while ATP was stable for the same period in freeze-dried samples. A strong metabolic stress caused by capturing the animals was evidenced by a marked lowering of the energy charge [(ATP)+1/2(ADP)/(ATP)+(ADP)+(AMP)], the natural level of energy charge being restored in about 24 h. The concentration of ATP showed no significant correlation with the dry weight of the individuals of a species nor with the mean dry weight of the species. The ATP concentration showed marked seasonal changes with increasing values during periods of reproduction, which is assumed to reflect an inheretn seasonal rhythm in the metabolism. ATP cannot be used as a general estimator of metabolism as it did not reflect differences in metabolic activity between different-sized individuals or species. The mean energy charge for 13 species was 0.7.Contribution from the Biological Station, University of Bergen, N-5065 Blomsterdalen, Norway.     

Grazing rates of the copepods Calanus glacialis and C. finmarchicus in arctic waters of the Barents Sea

Natural feeding rates of Copepodite Stages IV and V, and adult female Calanus glacialis (Jaschnov) and Copepodite Stage V and adult female C. finmarchicus (Gunnerus) were estimated using fluorescence analysis of gut contents. Measurements were made on copepods sampled from arctic waters east of Svalbard (Barents Sea) during the spring phytoplankton increase, in the period from 27 May to 13 June, 1983. Observations on Copepodite Stages IV and V and adult female C. glacialis suggest that the gastric evacuation rate is independent of developmental stage, whereas C. finmarchicus Copepodite Stage V showed a lower gastric evacuation rate than adult females. Gut fullness displayed a low correlation with the ambient chlorophyll concentrations. Ingestion rates calculated for C. glacialis were 0.3, 2.3, and 11.0 g C h^-1 for Copepodite Stages IV and V and adult females, respectively. Copepodite Stage V and adult female C. finmarchicus ingested 0.9 and 1.1 g C h^-1 at a temperature of ca.-1.0°C. The maximum ingestion rate in terms of percent body carbon d^-1 was higher for adult female C. glacialis and C. finmarchicus than for the respective Copepodite Stage V's. The results are discussed both in relation to the physiological state of the species and to the environmental conditions.     

Elemental and biochemical composition of salps (Tunicata: Thaliacea)

Carbon and nitrogen content have been measured in the solitary and aggregate generations of 11 species of salps. Regression equations for each species and generation permit estimation of carbon or nitrogen content as a function of length of live individuals. Different species of the same length may have nearly tenfold differences in carbon content. Fractionation and biochemical analysis of some samples revealed that the organic content of salps is approximately 80% protein. Ash-free dry weights average 27% of dry weights; mean carbon content is 29% of ash-free dry weight. Excess ash-free dry weight not accountable as organic material is thought to be water of hydration.     

Nitrogen limitation of Codium fragile ssp. tomentosoides as determined by tissue analysis

The critical nitrogen concentration of Codium fragile ssp. tomentosoides (van Goor) Silva was 1.90% on a dry weight basis. Internal nitrogen levels of thalli collected from three Rhode Island populations ranged from 0.75±0.08 to 3.72±0.08%. Internal nitrogen content was minimal in summer and maximal in winter, indicating nitrogen limitation during the summer and nitrogen storage during the winter. Part of this nitrogen storage appeared to support new growth of C. fragile in the spring. Carbon:nitrogen (C:N) ratios were inversely related to the internal nitrogen concentration. Low internal nitrogen content indicated nitrogen limitation better than C:N ratios because the C:N ratio could be influenced by both carbon and nitrogen metabolism.     

Respiration and excretion rates of Calanus glacialis in arctic waters of the Barents Sea

The respiration and excretion rates of Calanus glacialis (Jaschnov) Copepodite Stages III, IV, V, and adult females from the drift-ice area east of Svalbard (Barents Sea) were measured in shipboard experiments during the period from 27 May to 13 June, 1983. The phytoplankton biomass and abundance varied considerably between localities, but these variations were not generally reflected in the respiration and excretion rates of the copepod. The respiration and excretion rates of C. glacialis at the ambient temperature of-1.8°C (average respiration rates of 0.95, 0.73, 0.57, and 0.60 l O₂ mg^-1 dry wt h^-1 for Copepodite Stage III, IV, V, and adult females, respectively) were similar to those previously reported for other large-sized copepods from cold or temperate areas. Average respiration and excretion rates tended to decrease with incubation time or time after capture. Measurements on ten occasions within a period of 27 h after capture revealed excretion rates of ammonium ranging between 2.9 and 16.8 for C III, 3.7 and 21.1 for C IV, 1.3 and 28.4 for C V, and 1.6 and 18.7 for adult females, all expressed as nmol mg^-1 dry wt h^-1. In all experiments, excretion rates of inorganic phosphate varied between 0.7 and 1.5 (C III), 0.5 and 1.1 (C IV), 0.2 and 0.8 (C V), and 0.3 and 1.0 (adult females) nmol mg^-1 dry wt h^-1. Ratios of O:N, O:P, and N:P indicated that much of the metabolic energy was derived from catabolism of proteins. Comparison of the turnover rate of carbon and nitrogen showed, however, that nitrogen turnover was between 2.6 and 8.9 times higher than that of carbon. This may indicate that the copepods deaminate ingested protein, with the carbon skeleton of the amino acids subsequently being used in the synthesis of lipid compounds, possibly wax esters.     

Elemental composition (C,N, H) and energy in the development of Pagurus bernhardus (Decapoda: Pagurida) megalopa

No differences in development time and mortality were detected between starved and fed laboratory raised megalopa of Pagurus bernhardus. The average time of development in 138 megalopa was determined as 7.3±0.1 (95% CI) days. During megalopa development P. bernhardus loses about 7% in dry weight (DW), 17% in carbon (C), 6% in nitrogen (N) and 17% in hydrogen (H). During development C/N ratio and individual energy content descend about 14 and 22% respectively. Weight specific energy content decreases by 17% in the first 3 d and remains constant at 12.3±0.3 (95% CI) J·(mg DW)^-1 thereafter. About another 25% in individual energy content was lost by molting to crabs. The measured compounds do not follow a steady decrease. The possibility is discussed that a period of low energy cost (about the first half of development) alternates with times of higher energy expenditure mainly based on lipids. A fixed physiological program different from starvation capability is indicated for P. bernhardus megalopae. By comparing megalopae hatched in two different seasons and years reference is given to the variability in growth pattern.     

Digestion rates and assimilation efficiencies of siphonophores fed zooplankton prey

Digestion times and assimilation efficiencies are critical factors used in calculations of carbon and nitrogen budgets. Digestion times of natural copepod prey differed significantly among 4 genera of siphonophores (P>0.001), from a minimum of 1.6 h to a maximum of 9.6 h. Assimilation efficiencies, in contrast, were uniformly high; 87 to 94% for carbon and 90 to 96% for nitrogen. Nitrogen assimilation was consistently greater than carbon assimilation. Assimilation efficiencies calculated according to dry weight substantially underestimated assimilation of carbon and nitrogen, while calculations using ash-free dry weights and ash-free to dry weight ratios approached values for carbon assimilation. These values are appreciably higher than most of the assimilation efficiencies previously measured for a few other planktonic carnivores. These results indicate very efficient digestion of food by siphonophores in oceanic environments where prey densities are low.     

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.     

Weight and chemical composition of some important oceanic zooplankton in the North Pacific Ocean

Wet and dry weight, total carbon, nitrogen, hydrogen, and ash contents were determined on 33 species of zooplankton distributed predominantly in the open sea region of the North Pacific. Sampling covered the waters from 44°N to the equator. Average percentage of dry weight to wet weight was about 19% of all samples from the whole area. Percentage dry weight of carbon in copepods was on an average 51.5%. The highest value, 66.6%, was obtained in eggs of the copepod Pareuchaeta sarsi. Mixed zooplankton was assumed to contain carbon comprising about 35 to 45% of the dry weight. Carbon contained in the zooplankton biomass existing in the upper 200 m in the western parts of the northern North Pacific and Bering Sea during spring and summer was estimated to range from 20 to 85 mg C/m³. Nitrogen content varied considerably with localities. Average ratio of carbon to nitrogen was 8.5 in subarctic copepods, and 4.1 in subtropic-tropic copepods. This ratio also varied with season. In the copepod Calanus cristatus the ratio was highest (10.0) in May, immediately after the spring bloom of phytoplankton, when the animals contained much fat. The ratio fell to 5.1 in December. There seemed to be a large seasonal variation in boreal zooplankton due to great fluctuations of environmental conditions, especially the amount of food available; in tropical species the range was small because of environmental uniformity. Average hydrogen content was about 6 to 10%. The percentage of ash to dry weight amounted to 39.3% in pteropods and 3.4% in copepods.     

Reproductive biology of the common shrimp Crangon crangon (Decapoda: Crangonidae) in the central Irish Sea

Aspects of the reproductive biology of the common shrimp Crangon crangon (L.) were studied in Port Erin Bay, Isle of Man, Irish Sea. Size at sexual maturity was determined from the proportions of ovigerous females and of females with maturing ovaries. The size at which 50% of females are mature is estimated (±95% confidence intervals) as 12.5±0.48 mm carapace length. Based on the proportions of ovigerous females and of mature females, the main breeding season was from January to June. Mean ovarian dry weights indicated two broods (winter and summer), with females bearing winter broods (WB) having higher gonad indices than those with summer broods (SB). WB females with non-eyed eggs and with eyed eggs differed in the regression of ovarian dry weight on carapace length, indicating preparation for laying a second brood. In both broods the moult stages of berried females were related to egg stage. Moulting will occur following the release of the brood. During embryonic development, mean egg length and egg volume were larger in all stages, and the mean dry weight of individual eggs of all stages heavier, in WB than in SB; there was no difference in egg number, however. Consequently, reproductive investment, the proportion of female weight devoted to egg production, was 67% higher in WB (0.20±0.04) than in SB (0.12±0.03). There is a significant effect of egg volume on brood weight, but not on egg number. In both broods, egg number was a negatively allometric function of female body size in non-eyed eggs and an isometric function of female body size in eyed eggs. Brood mortality during incubation was higher in SB (17%) than WB (10%). Differences in the reproductive variables and investment between the two broods of C. crangon are discussed in the light of reproductive strategies and life history.Communicated by J.P. Thorpe, Port Erin     

Chemical changes during growth and starvation of larval Pleuronectes platessa

Plaice (Pleuronectes platessa L.) were sampled during periods of growth and starvation, from the end of the yolk-sac stage through metamorphosis, for changes in water, triglyceride, carbohydrate, total nitrogen, total carbon, and ash. The percentage of water in larvae decreased continuously during development. During post-hatching growth (up to late Stage 2) nitrogen and carbohydrate were laid down faster than triglyceride. The pattern changed during later larval development. The early deposition of protein in preference to neutral fat suggests that conversion of food during growth, without simultaneously laying down fatty energy stores, may be advantageous to pelagic marine fish larvae. During starvation the percentage of water in plaice larvae increased. Triglyceride, carbohydrate, nitrogen and carbon (as a percentage of the dry body weight) decreased during starvation, but ash increased sharply. The continuous use of nitrogen during starvation may be a catabolic adaptation to the marine environment.     

Microhabitat utilisation patterns in cryptobenthic coral reef fish communities

The cryptobenthic reef fish communities from four microhabitats at Orpheus Island, central Great Barrier Reef are described. Eighty-four 0.4m² samples yielded a total of 368 individuals from 42 species in eight families, with a mean density of 11 individuals m^–2 (±1.7SE) and 2.9 species 0.4 m^–2 (±0.2SE). Caves contained the highest number of both individuals (120) and species (26), followed by sand/rubble, soft coral, and open reefs. Microhabitat associations included cave and soft coral specialists. Site fidelity in 71 tagged individuals of 4 species was high, with a mean recapture rate of 53% (±8.4SE) remaining within the ~0.4 m² sampling area after a 48-h period. Behavioural observations also reflect this limited movement, with the dominant mode of behaviour in 7 species being a motionless state (67.5% ±11.6SE), followed by feeding (21.8% ±8.7SE), hiding (6.3% ±1.6SE), and swimming (4.4% ±1.5SE). Two distinct behavioural groups are identified: (1) sedentary forms, characterised by long periods of immobility (5 species); and (2) winnowers, characterised by long feeding bouts (2 species). The fine-scale partitioning of microhabitats, restricted home ranges, and sedentary behaviour of many cryptobenthic reef fish species suggest that this reef fish community exhibits similar patterns of habitat utilisation to their larger reef-fish counterparts, but at a much finer scale.     

Growth, carbon, nitrogen and caloric content of Solea senegalensis (Pisces: Soleidae) from egg fertilization to metamorphosis

Eggs and larvae of the Senegal sole, Solea senegalensis Kaup, were reared from fertilization until the end of metamorphosis, which occurs by Day 17 after hatching at 19.5 °C. Changes in energy content and biomass quality were studied in terms of dry weight and of carbon, nitrogen and energy content. S. senegalensis spawned eggs of about 1 mm diameter which hatched 38 h after fertilization. Average dry weight of individual eggs was 46 μg, the chorion accounting for about 18% of total dry weight. Gross energy of recently fertilized sole eggs was approximately 1 J egg⁻¹. From fertilization to hatching, eggs lost 8% of their total energy (chorion not included). After hatching, larvae lost 14% of their initial energy until the start of feeding which occurred about 48 h afterwards. The principal components catabolized during embryogenesis were carbon-rich compounds that decreased by 26%, while nitrogen-rich compounds decreased by only 10% and were practically unaltered from hatching to the start of feeding. Feeding larvae displayed constant growth during the period studied (specific growth rate on a dry weight basis was 0.26 d⁻¹). The relative proportion of carbon and nitrogen content revealed an accumulation of high energy compounds in the days before metamorphosis. By Day 14, the energy content reached values similar to those of recently hatched embryos, but decreased again during metamorphosis. Received: 10 June 1998 / Accepted: 28 January 1999     

Accumulation and loss of biomass inLiocarcinus holsatus larvae during growth and exuviation

Liocarcinus holsatus (Fabricius) larvae, of females originating from the Elbe Estuary, FRG, were reared in the laboratory at constant 15°C in May 1988. For each larval stage, developmental time was measured by individual cultures (Zoea I: 6.7±0.7d; Zoea II: 5.0±0.6d; Zoea III: 4.8±0.7 d; Zoea IV: 5.3±0.6d; Zoea V: 6.1±1.1d; Megalopa: 10.45±0.7d). During the entire period of development, dry weight (W), carbon (C), nitrogen (N), and hydrogen (H) were measured daily (Zoea I to V) or every second day (Megalopa). The energy content (E) was estimated from C. Biomass and energy (per individual) increased in each larval stage as a parabolic function of age and is described by power functions. C, H, and E exhibit a higher percentage gain (relative to initial values at the time of hatching) than W and N. It is suggested that proportionally more lipid than protein is accumulated during larval development. Cyclical changes in the relative biomass (% W) correspond to the larval moult cycle, indicating a rapid uptake of water and minerals immediately after hatching and a later increase in tissue growth. Changes in the C:N ratio suggest that during the first period more lipid than protein is accumulated. These patterns of growth and elemental composition are compared with literature data and a high degree of similarity in the growth characteristics of decapod larvae is seen. In addition W, C, N, and H values as well as E were measured for the exuviae of Zoea I to V and Megalopa. The percentage loss of growth rate by exuviae for each larval instar were higher in W (12 to 16%) and C (8 to 12%), and varied between 5 and 10% for N, H, and E.     

Respiration and excretion by the ctenophore Mnepiopsis leidyi

Respiration (dissolved oxygen and carbon dioxide) and excretion (dissolved organic carbon, inorganic and organic nitrogen and phosphorus) rates were measured for a variety of sizes of Mnemiopsis leidyi over a temperature range of 10.3° to 24.5°C. Both respiration and excretion rates were a direct linear function of animal weight and very temperature sensitive (Q₁₀4). Oxygen uptake ranged from 155 to 489 g at O/(g dry weight) day^-1 and carbon dioxide release from 43 to 166 M. Organic carbon made up about 38% of the total carbon released. Inorganic nitrogen excretion, exclusively in the form of ammonium, comprised 54% of the total nitrogen release and ranged from 10 to 36 M NH₄/(g dry weight) day^-1. Average release of dissolved primary amines (expressed as glycine equivalents) equaled 43% of the organic nitrogen fraction. Inorganic phosphorus release ranged from 2.0 to 4.9 M/(g dry weight) day^-1 and made up about 72% of the total phosphorus loss. The turnover of elements in the body was calculated as 5 to 19% per day for carbon and nitrogen, depending on the temperature, and an even higher 20 to 48% per day for phosphorus. These values are comparable to rates observed for small, active zooplankton.