Influence of food quantity and temperature on development and growth of the marine copepod Calanus chilensis from northern Chile

An experiment under laboratory conditions was conducted to test the hypothesis that development and growth of copepodite stages in Calanus chilensis are temperature-dependent and not subject to food shortage in the upwelling area of the Humboldt Current, northern Chile. Field data obtained from June 1994 to May 1995 in Bahía Mejillones (23°S) were used to define four combinations of temperature and food under which copepodites were reared from Stage CIII to adulthood. The high temperature was 18.1 °C and the low temperature 13.1 °C, whereas the high food level was in the range of 6.8 to 24.8 μg l⁻¹ chlorophyll a and the low level 1.0 to 6.8 μg l⁻¹ chlorophyll a. As food a mixture of three unknown species of phytoflagellates and the diatom Navicula cryptocephala was used. This phytoplankton was initially obtained from the same sampling sites as copepods and kept in f/2 media at stable levels and composition throughout the experiment. The development rate (1/t), estimated from the time (t) elapsing between Stage CIV and adult, was significantly affected by both temperature and food, although low-food effects were much more remarkable. Low-food conditions also significantly reduced body length and “structural” (lipid-discounted) body mass at adulthood, while temperature only affected body length. The weight-specific growth rate was also affected by food and temperature, but again food effects were much more drastic. The results indicate that C. chilensis is a highly sensitive species to lack of food, and is possibly subject to food shortage during its annual cycle in the coastal upwelling area of northern Chile. Food limitation may help explain the seasonal pattern of adult size reported by previous studies in the area and the lack of consistence between the number of generations predictable from a temperature-dependent model and that observed in the field during the annual cycle. Received: 10 September 1996 / Accepted: 29 October 1996     

Physioecology of zooplankton. IV. Effects of phytoplankton concentration,temperature, and body size on the net production efficiency ofCalanus pacificus

Weight-specific rates of individual production, total metabolic expenditure and assimilation, and net production efficiencies were estimated forCalanus pacificus Brodsky of selected body weights cultured at various phytoplankton concentrations and temperatures. The weight-specific rate of individual production increased hyperbolically with food concentration, and the maximum rate of individual production decreased logarithmically with a linear increase in body weight propotionally more at high than at low temperature. The weight-specific rate of total metabolic expenditure decreased logarithmically with increasing body weight and was unaffected by changes in food concentration. The effects of food concentration and temperature on the weight-specific rate of assimilation were similar to those on the rate of individual production, but the effect of body size differed considerably. The diversity in the temperature and body-size dependence of the maximum weight-specific rates of various physiological processes suggest (1) that, except for the metabolic rate, the allometric model (log-log relation) is inadequate for describing relationships between maximum rates of physiological processes and body size within species, and (2) that the common assumption that temperature affects the rates of various physiological processes in similar ways is not justified. Net production efficiency increased hyperbolically with food concentration, and the maximum production efficiency first increased slightly and then decreased gradually with increasing body weight. Small copepods attained higher efficiency at high temperature but larger ones did so at low temperature. The critical food concentrations for production efficiency and for the rate of individual production increased with increasing temperature and body size. Because of the effects of interactions among critical food concentration, temperature, and body size on the rates of growth and individual production and on net production efficiency, early development stages ofC. pacificus optimized growth and food conversion efficiency at high temperature, but late stages, particularly at low food concentrations, grew best and transformed food more efficiently at low temperature.Contribution No. 1130 From the Department of Oceanography, University of Washington, Seattle, Washington 98195, USA     

Growth and development of nauplii and copepodites of the estuarine copepod Acartia tonsa from southern Europe (Ria de Aveiro, Portugal) under saturating food conditions

A temperature-dependent growth model is presented for nauplii and copepodites of the estuarine calanoid copepod Acartia tonsa from southern Europe (Portugal). Development was followed from egg to adult in the laboratory at four temperatures (10, 15, 18 and 22°C) and under saturating food conditions (>1,000 μg C l⁻¹). Development times versus incubation temperature were fitted to a Belehradek’s function, showing that development times decreased with increasing incubation temperature: at 10°C, A. tonsa need 40.3 days to reach adult stage, decreasing to 8.9 days when reared at 22°C. ANCOVA (homogeneity of slopes) showed that temperature (P<0.001) and growth phase (P<0.01) had a significant effect on the growth rate. Over the range of temperatures tested in this study, highest weight-specific growth rates were found during naupliar development (NI–NVI) and varied from 0.185 day⁻¹ (10°C) to 0.880 day⁻¹ (22°C) with a Q ₁₀ equal to 3.66. During copepodite growth (CI–CV), the weight-specific growth rates ranged from 0.125 day⁻¹ (10°C) to 0.488 day⁻¹ (22°C) with a Q ₁₀ equal to 3.12. The weight-specific growth rates (g) followed temperature (T) by a linear relationship and described as ln g=−2.962+0.130 T (r ²=0.99, P<0.001) for naupliar stages and ln g=−3.134+0.114T (r ²=0.97, P<0.001) for copepodite stages. By comparing in situ growth rates (juvenile growth and fecundity) for A. tonsa taken from the literature with the temperature-dependent growth model defined here we suggest that the adult females of A. tonsa are more frequently food limited than juveniles.     

Importance of food quantity to structural growth rate and neutral lipid reserves accumulated in Calanus finmarchicus

 Growth and developmental rates were determined for copepodids of Calanus finmarchicus (Gunnerus) from experimental seawater mesocosms in a western Norwegian fjord. The instantaneous growth rates (g) from copepodid stage I (CI) to adult ranged from 0.08 to 0.10 d⁻¹. Daily per capita mortality rate of the cohorts was as low as 0.012 d⁻¹ (1.2% d⁻¹). At local increasing temperatures (5.1 to 8.3 °C), development was equiproportional, and the cumulative median development time from egg to CV was approximately 65 d. CV moulted to males and females, and egg production was initiated. Enhancement of food resources by nutrient addition caused a 23.4% increase in growth rates from CI to adult. Additionally, copepodid stages showed a generally larger body size, carbon and nitrogen content and total storage lipid content (wax esters + triacylglycerols) in response to enhanced resources. Our data support an elsewhere proposed exponential-growth hypothesis; growth of the structural compartments and store lipids (mostly wax esters) was exponential during the copepodid stages. However, a sigmoidal pattern of growth best described growth of adult stages if reared at high resources, and depot lipid accumulation in late CVs and adults at high resources. Body nitrogen growth increased exponentially, however, no significant changes in nitrogen specific growth rates were found between individuals from low and high resources. CV and adults seem to have reached near-maximal weights at high resources, whereas structural weight continued to increase at low resources. Despite the differences in structural growth dynamics, cohort development was similar until the end of CV. During the onset of sexual differentiation, the male:female ratio and the adult:CV ratio were highest at high food resources, suggesting that the time used for the final moult depends on the feeding history of the copepods in relation to food quality and quantity. It appears that relatively small changes in food availability strongly influence the biochemical composition of C. finmarchicus copepodids. Received: 28 May 1999 / Accepted: 10 February 2000     

Physioecology of zooplankton. I. Effects of phytoplankton concentration,temperature, and body size on the growth rate of Calanus pacificus and Pseudocalanus sp.

Changes in dry weight and in weight-specific growth rates were measured for copepodite stages of Calanus pacificus Brodsky and Pseudocalanus sp. cultured under various combinations of phytoplankton concentration and temperature. Mean dry weight of early copepodites was relatively unaffected by either food concentration or temperature, but mean dry weight of late stages increased hyperbolically with food concentration and was inversely related to temperature. The food concentration at which maximum body weight was attained increased with increasing temperature and body size, and it was considerably higher for C. pacificus than for Pseudocalanus sp. This suggests that final body size of small species of copepods may be determined primarily by temperature, whereas final body size of large species may be more dependent on food concentration than on temperature. Individual body weight increased sigmoidally with age. The weight-specific growth rate increased hyperbolically with food concentration. The maximum growth rate decreased logarithmically with a linear increase in body weight, and the slope of the lines was proportional to temperature. The critical food concentration for growth increased with body size proportionally more at high than at low temperature, and it was considerably higher for C. pacificus than for Pseudocalanus sp. Because of these interactions, early copepodites optimized growth at high temperature, even at low food concentrations, but under similar food conditions late stages attained higher growth at low temperature. The same growth patterns were found for both species, but the rates were significantly higher for the larger species, C. pacificus, than for the smaller one, Pseudocalanus sp. On the basis of findings in this study and of analyses of relationships between the maximum growth rate, body size, and temperature from other studies it is postulated (1) that the extrapolation of growth rates from one species to another on the basis of similarity in body size is not justified, even for taxonomically related species; (2) that the allometric model is inadequate for describing the relationship between the maximum weight-specific growth rate and body size at the intraspecific level; (3) that the body-size dependence of this rate is strongly influenced by temperature; and (4) that species of zooplankton seem to be geographically and vertically distributed, in relation to body size and food availability, to optimize growth rates at various stages of their life cycles.Contribution No. 1127 from the Department of Oceanography, University of Washington, Seattle, Washington 98195, USA     

Relationship between growth and biochemical indices in laboratory-reared juvenile Japanese flounder (Paralichthys olivaceus), and its application to wild fish

In order to estimate growth rates based on biochemical indices of the liver of wild Japanese flounder (Paralichthys olivaceus), juveniles were reared at six ration levels (0, 0.5, 2, 4, 6 and 8% body weight day⁻¹) in the laboratory for 14 days, and the relationship between their growth rates and biochemical indices (RNA/DNA, protein/DNA, triglyceride/DNA, phospholipid/DNA and cathepsin D activities) were determined. Positive and approximately linear relationships were seen between growth rates and the indices of RNA/DNA, protein/DNA and phospholipid/DNA. The triglyceride/DNA ratio decreased with increasing growth rates up to approximately 1% body weight day⁻¹, then increased linearly with increasing growth rates. There was no significant correlation between growth rates and cathepsin D activity, and the highest values were obtained in the starved fish. Compared with laboratory-reared specimens, wild specimens of similar sizes were found to have significantly larger livers. The RNA/DNA, protein/DNA and phospholipid/DNA ratios of wild specimens fell in a broad range between ration groups of reared juveniles. The protein/DNA ratios of wild specimens were low and outside the range of the reared juveniles at six ration levels. In contrast, the levels of cathepsin D activity of wild fish were highest compared to the reared fish. Estimated growth rates of wild fish from the RNA/DNA, protein/DNA and phospholipid/DNA regressions obtained from the rearing experiment were 1.66, −1.74 and 0.10% day⁻¹, respectively. Based on our results, the RNA/DNA index may be regarded as the most valid and reliable growth estimator. It is noted that the larger liver size, the lower liver protein/DNA ratio and the unexpectedly high level of cathepsin D activities of wild specimens found in this study may reflect the different metabolic conditions of fish reared in the laboratory compared to those collected in the field. Received: 29 February 2000 / Accepted: 26 August 2000     

Zooplankton growth rates: the influence of size in nauplii of tropical marine copepods

Growth rates and development times were determined for nauplii of the genera: Acartia, Centropages, Corycaeus, Oithona, Paracalanus, Parvocalanus and Temora in nearshore waters of Jamaica from in situ microcosm incubations. At these high local temperatures (∼28 °C), total naupliar development time was short: 3 to 4 d inshore and 4 to 5 d offshore. Mean instantaneous growth rates (g) ranged from as high as 0.90 d⁻¹ for Parvocalanuscrassirostris to as low as 0.41 d⁻¹ for Corycaeus spp. In general, nauplii of cyclopoid copepods appeared to grow more slowly than those of calanoids of the same size. Naupliar growth rates were significantly related to body size (r ²= 0.43 to 0.50), but were unrelated to chlorophyll concentration in any measured size-fraction. This suggests that nauplii are generally not limited by resources, but are growing at their maximum temperature and size-dependent rates. Received: 30 May 1997 / Accepted: 13 May 1998     

Impact of temperature on juvenile growth and age at first egg-laying of the Pacific reef squid Sepioteuthis lessoniana reared in captivity

Cephalopod mollusks exhibit highly plastic life cycle traits influenced primarily by the interactive effects of food availability, light cycle and temperature, with the latter perhaps the most influential. Hatchlings of the tropical reef squid Sepioteuthis lessoniana were hatched from field-collected eggs in the laboratory and cultured at different temperatures to evaluate the effect of temperature on growth rates. All groups showed rapid, sustained growth rates from hatching to a size of 10–25 g. Beyond this size range, growth was slower and not clearly exponential in form. Growth rate was closely linked to temperature. Squids grown at approximately 27 °C attained a size of 10 g in as little as 45 days at sustained growth rates of 12.2% body weight day⁻¹ (%bw day⁻¹), while squids cultured at 20 °C required almost 100 days to attain the same size at rates of 5.7%bw day⁻¹. At an age of 55 days and approximately 1 g body weight, juvenile squids cultured at 20 °C were able to accelerate growth rates from 5.7%bw day⁻¹ to over 12%bw day⁻¹ when temperature was raised to 27 °C. They maintained this growth rate to a size of about 10 g and an age of at least 75 days post-hatching, indicating that body size and not age is the limiting factor for this rapid post-hatching growth. By comparison, conspecifics cultured near 27 °C from hatching had shifted out of the rapid post-hatching growth phase by day 50 at sizes between 10 and 50 g. The hatchlings from temperate to subtropical Japan had consistently higher growth rates at comparable temperatures than hatchlings from tropical Okinawa. When plotted as growth rate versus temperature, the Japanese group had a clearly higher slope to the relationship than the tropical populations, equivalent to a 2%bw day⁻¹ difference in growth rate at 25 °C. Age at first egg-laying was decreased at higher culture temperatures; however, overall life span was not. Received: 21 February 2000 / Accepted: 6 September 2000     

<Emphasis Type="Italic">Oithona similis</Emphasis> in a high latitude ecosystem: abundance,distribution and temperature limitation of fecundity rates in a sac spawning copepod

In this study we report the abundance, fecundity and an index of the relative survival of Oithona similis (nauplii to copepodites) across a large latitudinal and temperature range within the Southern Ocean. The abundance of O. similis was strongly related to temperature and to depth-integrated (0–100 m) chlorophyll a, abundance increasing with increasing temperature (and therefore decreasing latitude) and Chl a. In situ egg production rates and fecundity per female were also significantly and positively related to temperature and Chl a. Egg hatch times rapidly lengthen as temperature decreases and in sac spawning species the next batch of eggs cannot be produced until the previous clutch hatch. Consequently, we predict that in O. similis, fecundity must decline rapidly at low temperatures, especially below 5°C. A model comparing maximum rates of fecundity with in situ data suggested production rates were strongly food limited across our study region. However, the relationship of in situ and maximum rates to temperature were similar, confirming the importance of temperature. Further, as time taken to develop from egg to adult also rapidly extends with declining temperature, it is increasingly unlikely that O. similis will be able to maintain its population against mortality. Our findings have broad implications for the cold temperature (and hence geographic) limits of O. similis, but also for the distribution of other sac spawning copepods and planktonic species generally.     

Growth and development rates of Calanus finmarchicus nauplii during a diatom spring bloom

Growth and development rates were determined for nauplii of Calanus finmarchicus (Gunnerus) in the near-shore waters of a western Norwegian fjord from in situ mesocosm incubations. The major food source for the nauplii was diatoms, but Phaeocystis sp., dinoflagellates and ciliates were also part of the diet. At local temperatures ranging from 4.8 to 5.2 °C the cumulative median development time from hatching to Nauplius VI was 19 d. The time taken to molt to the next naupliar stage was approximately constant (3 d) from Stages IV to VI, but Stage III needed the longest development time (5 d). The instantaneous growth rate in terms of body carbon was negative from hatching to Nauplius Stage II, but as high as 0.25 to 0.30 d⁻¹ from Stage III to V. Enhancement of food resources by nutrient addition led to no significant change in specific growth rates. Additionally, the cohorts from different nutrient regimes showed almost equal development time, size and body carbon within stages. Length–weight relationships of nauplii from the two different food resources were: W _{low resources} = 4.17 × 10⁻⁶ × L ^2.03 (r ² = 0.84) and W _{high resources} = 4.29 × 10⁻⁶ × L ^2.05 (r ² = 0.92), where weight (W) is in micrograms of C and body length (L) in micrometers. The natural body morphology of naupliar stages I to VI is illustrated with digital images, including the final molt from Nauplius VI to Copepodid Stage I. In general, development of the nauplii was faster than that of the copepodids of C. finmarchicus, and structural growth was exponential from naupliar stages III to VI. This study validates our earlier results that nauplii of C. finmarchicus can obtain high growth and nearly maximal developmental rates at relatively low food levels (∼50 μg C l⁻¹), suggesting that nauplii exhibit far less dependence on food supply than copepodids. Received: 30 July 1999 / Accepted: 7 March 2000     

Zooplankton growth rates: the influence of size and resources in tropical marine copepodites

Growth rates were determined for copepodites of the genera: Acartia, Centropages, Corycaeus, Oithona, Paracalanus, Parvocalanus and Temora in nearshore waters of Jamaica from in situ microcosm incubations. At these high local temperatures (∼28 °C), total copepodite development time was as short as 4 to 5 d. Mean instantaneous growth rates (g) ranged from as high as 1.2 d⁻¹ to as low as 0.1 d⁻¹. In general, cyclopoid copepods appeared to grow more slowly than calanoids of the same size. Enhancement of resources by nutrient addition caused a 32% increase in growth rates in experiments from a mesotrophic site, but only a 17% increase at a more eutrophic site. Additionally, copepodites at both sites showed faster development and generally larger size at stage in response to nutrient addition. Growth rates were positively related to chlorophyll concentration in the >2 μm size-fraction. A significant relationship of growth rate to body size (r ² = 0.45) emerged across a wide range of trophic status, but it was confounded with resource availability. It appears that growth in tropical copepod copepodites may be frequently limited by resources in a size-dependent manner. Received: 30 May 1997 / Accepted: 13 May 1998     

Spectral properties and photosynthetic action in red-tide populations of Prorocentrum micans and Gonyaulax polyedra

Three sets of experiments were carried out between April and September 1987 to assess reproductive energetics in a laboratory population of the capitellid polychaete Capitella capitata type I. In the first experiment, the population was sampled intensively over a short period of 3 wk. Weight-specific reproductive output did not differ significantly between females and hermaphrodites. However, because hermaphrodites were significantly bigger than females, their fecundity was significantly higher. In the second experiment, fecundity and weight-specific reproductive output were monitored weekly over the oscillation cycle of the population (30 wk). Both fecundity and weight-specific reproductive output varied drastically (40 to 80 eggs per female and 17 to 57%, respectively). In the third experiment, non-sexed juveniles were reared for 15 d on six ration levels (normalized on the basis of organic nitrogen) of four different food sources. The maximum weight-specific reproductive output was 70% (for females). Both fecundity and weight-specific reproductive output were positively correlated with food ration and varied with food type. The data on weight-specific reproductive output recorded in the present study have been compared with those of other iteroparous polychaetes in the literature. Reproductive output in C. Capitata type I varies more widely and is greater than in other iteroparous polychaetes. In this species, reproductive output and population dynamics vary concomitently with the level of organic matter in the habitat.     

Metabolism and elemental composition of four oncaeid copepods in the western subarctic Pacific

Respiration rates and elemental composition (carbon and nitrogen) were determined for four dominant oncaeid copepods (Triconia borealis, Triconia canadensis, Oncaea grossa and Oncaea parila) from 0–1,000 m depth in the western subarctic Pacific. Across the four species of which dry weight (DW) varied from 2.0 to 32 μg, respiration rates measured at in situ temperature (3°C) increased with DW, ranging from 0.84 to 7.4 nl O₂ individual⁻¹ h⁻¹. Carbon (C) and nitrogen (N) composition of the four oncaeid species ranged from 49–57% of DW and 7.0–10.3% of DW, respectively, and the resultant C:N ratios were 4.8–8.3. The high C contents and C:N ratios were reflected by large accumulation of lipids in their body. Specific respiration rates (SR, a fraction of body C respired per day) ranged between 0.5 and 1.3% day⁻¹. Respiration rates adjusted to a body size of 1 mg body N (i.e. adjusted metabolic rates, AMR) of the four oncaeid species [0.6–1.1 μl O₂ (mg body N)^−0.8 h⁻¹ at 3°C] were significantly lower than those (1.7–5.1) reported in the literature for oithonid and calanoid copepods at the same temperature. The present results indicate that lower metabolic expenditure due to less active swimming (pseudopelagic life mode) together with rich energy reserve in the body (as lipids) are the characters of oncaeid copepods inhabiting in the epi- and mesopelagic zones of this region.     

Ontogenetic variation in metabolism, biochemical composition and energy content during the early life stages of Farfantepenaeus paulensis (Crustacea: Decapoda: Penaeidae)

Dry weight (DW), oxygen consumption, ammonia-N excretion, proximate biochemical composition (total protein, carbohydrate, lipid, water and ash), and energy content (estimated from biochemical composition and by wet combustion) were determined in early developmental stages of cultured Farfantepenaeus paulensis. Pooled samples from embryonic, larval and postlarval stages (at 26 ± 1 °C and 34 ± 1‰) were used for measurements. The study focused on physiological and biochemical processes during transitional periods of ontogeny, such as hatching, lecithotrophic and planktotrophic stages, metamorphosis, and the attainment of a benthic existence in postlarva. DW showed higher increment between protozoea I (PZ I) and mysis I (M I) than in the next mysid and postlarval stages. Individual rates of oxygen consumption and ammonia-N excretion increased, while weight-specific rates presented significant reduction throughout development. Higher weight-specific oxygen consumption was registered in nauplius III (N III) and PZ I, following a decrease in subsequent stages. Postlarval stages PL V–VI and PL X–XII exhibited the lowest values among the stages studied. Weight-specific excretion was high in N III and protozoeal stages, with maximum values in PZ II, while the following stages were marked by lower rates. O:N ratios indicated higher protein catabolism in the stages between egg and M I and a shift to more lipid utilization close to metamorphosis. Water content was higher in the protozoeal stages and decreased afterwards. Higher percentages of protein, lipid and carbohydrate (%DW) were observed in egg and nauplius stages. Protein and lipid decreased from the egg through the naupliar and protozoeal stages, rising again in mysis stages. Lipid content (%DW) decreased in PL V–VI and PL X–XII. Lipid:protein ratios showed an increase of the importance of lipid between PZ III and M II. Carbohydrates represented a minor fraction of body composition, and ash percentages increased from egg to a maximum in PZ II, decreasing in subsequent stages. Energy content determined by wet combustion or calculated by energy equivalents presented the same trend throughout development, varying similarly to protein. Protein was the main energy contributor to body energy in all stages, while the importance of lipid was higher in egg and early naupliar stages. Trends observed in metabolic rates and body composition may be associated to morphological and behavioral changes during the early stages of penaeid development, such as the transition from herbivory to omnivory, and the adoption of a benthic existence. Different ontogenetic energy strategies contribute to succeed through such diverse type of development. Received: 4 July 2000 / Accepted: 6 December 2000     

Sexual reproduction of the scyphomedusa Aurelia aurita in relation to temperature and variable food supply

The effects of food availability and temperature on sexual maturation and female reproductive output of the scyphomedusa Aurelia aurita was examined in two populations from the contrasting environments of Southampton Water and Horsea Lake, England. Trends in oogenesis and subsequent reproductive output differed markedly between the two populations. In Southampton Water, the onset of sexual maturation occurred earliest in the larger medusae, but eventually all females became ripe, the smallest being 45 mm bell diameter (BD). The decrease in minimum size at maturity was correlated with increasing temperature. In A. aurita from Horsea Lake, size at maturity varied on a seasonal basis, with the smallest ripe female being only 19 to 20 mm BD. There were spring and autumn periods of sexual maturation in this population. During the autumn period, it is likely that food limitation was playing a more critical role in determining medusa size, with decreasing temperature indirectly affecting A. aurita by limiting primary and secondary production. In similar-sized ripe medusae, fecundity was greater in Southampton Water, but the planula larvae produced were significantly smaller than those in Horsea Lake. It is suggested that in Horsea Lake, the quality of the larvae are greater in terms of biochemical content to ensure survival of the few gametes produced (i.e. K-strategy). Comparison of the reproductive effort of the two A. aurita populations revealed that medusae from Southampton Water, which experience greater food availability, are able to direct more energy to reproduction than Horsea Lake medusae. In the latter, A. aurita medusae appear to partition the available food resources into either somatic growth (and therefore increased future fecundity) when food is abundant, or reproductive growth when food is scarce. Received: 24 June 1997 / Accepted: 23 March 1998     

Bioenergetics of ephyra larvae of the scyphozoan jellyfish Aurelia aurita in relation to temperature and salinity

The body mass of Aurelia aurita ephyrae was better correlated with the diameter of the central disc than with the distance between opposite rhopaliae or distance between opposite lappet tips. Body dry weight (y, in μg) related to the disc diameter (x, in mm) through the equation y = 22.33 x ^1.99. The exponent 1.99 was significantly lower than that for the medusa stage, indicating a tendency to grow in diameter rather than in weight through the ephyra stage. The average ash-free dry weight (AFDW) of ephyrae was 38.0% of the dry weight. The AFDW/diameter relationship was used to convert measured diameters to body AFDW and calculate succession in body mass, daily ration, daily growth rate and gross growth efficiency. Effects of temperature (6, 9.5, 12, 15 and 18 °C) and salinity (17.5, 22, 26, 30.5 and 35 PSU) on these parameters and feeding were studied at saturated prey concentration (222 Artemia nauplii l⁻¹ initial concentration) by daily measurements over 10 d. There was a strong effect of temperature for total ingestion, growth rate, growth efficiency and final body mass of individual ephyrae, whereas the daily ration was not significantly different between the different temperatures. The experimental group kept at the highest temperature (18 °C) diverged the most, and ephyrae at this temperature ingested 2.7 times more and increased in weight 5.4 times more than at 6 °C. The average daily growth rate and gross growth efficiency of these ephyrae were 34.5% and 25.1%, respectively, significantly higher than at 6, 9.5 and 15 °C. Significant effects of salinity were shown for total ingestion, daily ration, daily growth rate and final weight, although only total ingestion and daily ration diverged sufficiently to show effects in a post-hoc test. This test showed that total ingestion was significantly different for all salinities except between 22 and 35 PSU and between 17.5 and 26 PSU. The daily ration for 35 PSU diverged from all other salinities, whereas none of the other salinities showed any significant differences. Thus, provided food in excess A. aurita can double its weight every 2 to 4 d, dependent on temperature and can therefore develop to the medusa stage in short time. Differences in environmental salinity in the range 17.5 to 35 PSU have little or no effect on growth rate and growth efficiency, whereas our results indicate that the full seawater salinity (35 PSU) causes significantly higher ingestion rate compared to lower salinities. Received: 11 January 1999 / Accepted: 11 May 1999     

Fecundity and spawning in a deep-water cirromorph octopus

 Reproductive characteristics of cirromorph octopuses, assigned to the species Opisthoteuthis grimaldii, were sampled as a commercial fishing by-catch on the Hebrides Slope, west of Scotland. A total of 254 specimens (99 female, 155 males), retrieved from bottom trawls fished at 750 to 1500 m depth, were examined. A maximum of 2097 eggs was counted in a single female ovary (mean female body weight 1242.8 g), most of them <1 mm in length. At egg lengths over 1 mm, diminishing numbers of eggs were present in 1 mm size categories up to a maximum of ≈10 mm. At body sizes >500 g (wet wt), and in every female >750 g (max. female weight recorded in the sample was 2959 g), a succession of unattached eggs was present in the proximal oviduct and a single, unattached mature egg occupied the tip of the distal oviduct. These females were assumed to be in spawning condition and the characteristics of egg distribution in the reproductive tract to be consistent with sequential release of individual eggs and continuous spawning throughout the growth period and lifespan of the mature octopus. In pre-spawning females there was a positive relationship between estimated egg numbers and maximum egg size. After the onset of spawning there was no significant further increase in estimated potential fecundity over the body-size range 500 to 3000 g. Follicular sheaths remaining in the ovary after release of eggs into the proximal oviduct were counted and used to estimate the total number of eggs released up to the time of capture. Follicular sheaths first appeared at 500 to 650 g body weight and increased steeply in number to >1000 in females >1500 g. Two individuals were found with ovarian follicular sheaths but with no terminal egg in the distal oviduct; these were assumed to have released their egg just before capture. Summation of the number of follicular sheaths counted plus the number of eggs estimated as remaining attached in the ovisac, provided a revised estimate of total potential fecundity and raised the estimate for any individual to a maximum of 3202 eggs (mean = 1396 eggs). Received: 3 February 2000 / Accepted: 17 May 2000     

Reproductive patterns of scleractinian corals in the northern Red Sea

The majority of published accounts on scleractinian coral reproduction are from the tropical Pacific and Caribbean, with very little information known about Red Sea species. This report examines variation in reproductive mode in 24 species of hermatypic corals (belonging to seven families) in the Gulf of Eilat, Red Sea. Eighteen species are hermaphroditic broadcasters, two are hermaphroditic brooders and three are gonochoric broadcasters. In the Pocilloporidae, the gonads project into the body cavity, while in the other six families the gonads reside inside the mesenteries. The number of gonads per polyp in broadcasting species follows family or genus lines. Fecundity (eggs per polyp) increases with polyp size. Brooding species usually exhibit one or two gonads per polyp and each gonad contains only one to three oocytes. Oocyte size varies widely and does not relate to mode of reproduction. The largest oocytes (diameter = 450 μm) occur in the brooding coral Alveopora daedalea and in broadcasting species of the genus Acropora (diameter = 420 μm). Gonad morphology and gonochorism versus hermaphroditism appear to be constrained phylogenetically at the family or genus level. Lastly, this report compares the data presented for Red Sea scleractinian species with the data available on scleractinian corals from other geographical regions. Received: 2 February 1993 / Accepted: 9 March 1998     

Physiological traits associated with a reproductive-resting stage in Coullana canadensis (Copepoda: Harpacticoida)

We investigated physiological traits associated with a reproductive-resting stage in adult female harpacticoid copepods, Coullana canadensis (Willey). Our hypothesis was this stage represents a life-history strategy that could increase fitness by improving winter survivorship and future reproductive success in spring. To test if physiological rates are suppressed in this stage, we compared gut-cell morphology, nitrogen excretion rates, enzyme activities, and phytoplankton grazing rates of reproductive-resting and reproductive females reared in the laboratory under high food conditions. Copepods came from laboratory cultures originating from individuals collected in Maine and Maryland, USA in 1990. Reproductive-resting females had lower physiological rates, and the surface area of gut-cells was reduced compared to reproductive females. Distinct morphological differences in the distribution of lipids between reproductive and reproductive-resting females were observed under light and electron microscopy, the latter having a diffuse accumulation of lipid in the area normally occupied by the ovaries. Differences in lipid composition were also found. Reproductive copepods had a significantly higher ratio (%) of polar lipids to total lipid, and a lower proportion of triacylglycerols compared to reproductive-resting copepods. These laboratory findings are consistent with the hypothesis that the reproductive-resting stage in C. canadensis is an adaptive response to increase winter survival. Received: 29 May 1996 / Accepted: 3 December 1997     

Egg production of Calanus finmarchicus : effect of temperature, food and season

The use of the egg production rate of herbivorous copepods as an important parameter for understanding population dynamics and as an index of secondary production requires knowledge of the regulatory mechanisms involved and of the response to changes in food concentrations and temperature. Furthermore, the effects of season and generation on egg production have to be studied. In this context data are presented for Calanus finmarchicus from the northern North Atlantic. Prefed and prestarved females were exposed to different concentrations of the diatom Thalassiosira antarctica over 1 to 2 wk at 0 or 5 °C, and egg deposition was controlled daily. Egg production increased with higher food concentrations, but much less when prestarved. The effect of temperatures between −1.5 and 8 °C on egg production was studied in females maintained at optimum feeding conditions. Egg production rate increased exponentially over the whole temperature range by a factor of 5.2, from 14.2 to 73.4 eggs female⁻¹ d⁻¹, and carbon-specific egg production by 4, from 2.1 to 8.5% body C d⁻¹. The response to starvation was also temperature dependent. In both the temperature and feeding experiments egg production rate was regulated mainly by changes of the spawning interval, while changes of clutch size were independent of experimental conditions. Different responses to optimum feeding conditions were observed in females collected in monthly intervals on three occasions between March and May. The March females deposited more clutches than the April and May females. In May, >50% of the females did not spawn at all. Maximum egg production rates were never >25% of the rate expected at 5 °C, indicating endogenous control of egg production in addition to food and temperature effects. Received: 4 August 1996 / Accepted: 11 September 1996