Interannual, seasonal, local and body size variations in reproduction of the prawn Penaeus (Marsupenaeus) japonicus (Crustacea: Decapoda: Penaeidae) in the Ariake Sea and Tachibana Bay, Japan

 The reproductive biology of female Penaeus japonicus Bate was investigated in the Ariake Sea and Tachibana Bay (located outside the Ariake Sea), Japan from 1994 to 1996. Interannual, seasonal, individual female body size and spatial influences on the incidence of spawning were examined. The proportion of inseminated females, increased with increasing body size up to 170 mm body length (BL) and decreased thereafter. The minimum size at maturity was similar between years; 130 to 140 mm BL. The minimum size of individuals with developing ovaries or spermatophores differed from that of ripe females by 20 to 25 mm. Spawning occurred in the central part of the Ariake Sea and Tachibana Bay but rarely in the inner part. P. japonicus had a clear reproductive cycle. Spawning started earlier and ended later, occurring from mid-May to mid-October, in the eastern central part of and outside of the Ariake Sea compared with the western central part, where spawning occurred from mid-June to mid-September. Spawning dynamics differed across seasons, body sizes and areas but not across years. The seasonal peak in the proportion of ripe females varied with body size. In small individuals (130 to 169 mm BL), no peak was observed, whereas in large individuals (>170 mm BL) the proportion of ripe individuals peaked in June. The proportion of ripe individuals increased with increasing body size and was high outside the Ariake Sea. Factors causing the variation in spawning dynamics are discussed. Received: 30 November 1998 / Accepted: 12 October 1999     

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     

Egg adoption can explain joint egg-laying in common eiders

Hypotheses regarding the evolution and maintenance of intraspecific nest parasitism were tested with data collected during a 3-year study of common eiders (Somateria mollissima) breeding near Churchill, Manitoba. The nest parasitism rate was highest (42.4% of nests) during the year with the highest nest density and the best environmental conditions, and lowest (20.2% of nests) in the year with the lowest nest density and the poorest environmental conditions. Over the nesting season, parasitic eggs were laid at the same time as normally laid eggs. Most parasitic eggs (>75%) were laid before the host female laid her third egg. The majority of the parasitic eggs were the first or second egg produced by the parasitic female. When a parasitic egg was laid before or on the same day as the host female initiated her clutch, the probability of her first egg being depredated before incubation was significantly lowered. First- and second-laid eggs suffered a high rate of predation probably because nesting females do not attend their clutch until their second or third egg is laid. Hypotheses that some females use intraspecific nest parasitism to parasitize the parental care of other females were inconsistent with these data. Egg adoption is a likely explanation for the prevalence of females incubating parasitic eggs in this population. Received: 30 September 1997 / Accepted after revision: 6 May 1998     

Life history of Gonatus onyx (Cephalopoda: Teuthoidea): deep-sea spawning and post-spawning egg care

 A reproductive strategy consisting of deep- water spawning and egg-care was inferred for the midwater squid Gonatus onyx Young, 1972. Brooding females and associated eggs and hatchlings, captured between 1250 and 1750 m depth off southern California, are described. Brooding females appear to be senescent and lack tentacles. Large eggs of this species (3 mm) at cold temperatures (3 °C at capture depth) may require as long as 9 mo to develop. The high lipid content of the digestive gland in adult females of this species may provide fuel to support such an extended “brooding” period. Received: 22 February 1999 / Accepted: 25 May 2000     

Fecundity of the chokka squid Loligo vulgaris reynaudii on the southeastern coast of South Africa

Potential fecundity in Loligo vulgaris reynaudii was estimated to be about 17 000 eggs, calculated as the total number of discernible oocytes in the ovary and oviduct. Squid were observed to spawn up to 8140 eggs over a 36 h period in captivity. First estimations of actual fecundity are therefore between 8000 and 17 000 eggs. Factors complicating a more accurate estimation of actual fecundity in this species include difficulties with aquarium maintenance, their behaviour of spawning over a protracted period and in multiple sites, and atretic oocytes observed in both developing and mature ovaries. Detailed morphological and histological analysis of gonads collected at regular intervals over a complete spawning season will allow a more precise calculation of actual fecundity. Received: 9 October 1998 / Accepted: 22 April 1999     

Flexible reproductive strategies in tropical and temperate Sepioteuthis squids

A major difficulty confronting the determination of cephalopod reproductive life history is assessing over what portion of the life span an individual is reproductively mature and actively depositing eggs. This paper assesses the potential of the tropical Sepioteuthis lessoniana and two genetic types of the temperate Sepioteuthis australis, to spawn multiple batches of eggs at discrete times throughout the adult life span. This is achieved by histological examination of the ovarian gametogenic cycle and detailed morphological assessments of the reproductive system, in conjunction with other biological information. The genetic type of S. australis found at the northern limits of its Australian distribution showed evidence of a high correlation between body size and quantity of mature eggs, suggesting that eggs may be accumulating to be laid in a single batch. Although maturation was also a size-related process in S. lessoniana and Tasmanian S. australis, oviduct size was not correlated with body weight in mature females, which is indicative of multiple spawning. Further supporting evidence includes relatively low gonadosomatic indices, the heavier weight of the ovary relative to the oviduct, and the feeding activity of mature animals. Mature S. lessoniana and S. australis individuals were present at each location over very wide age and size ranges. In Tasmanian waters, there were distinct seasonal differences in the reproductive biology of S. australis. Summer-caught individuals had much higher gonadosomatic indices and may have been laying larger batches of eggs compared with winter-caught individuals. Summer-caught females also showed a negative correlation between egg size and egg number within the oviduct, suggesting that some individuals were producing fewer, larger eggs and others many smaller eggs. Evidence suggests that considerable flexibility is inherent in the reproductive strategy of both S. lessoniana and S. australis. Received: 11 May 2000 / Accepted: 12 September 2000     

Reproductive biology of deep-water calanoid copepods from the Arctic Ocean

Reproductive modes and egg production were studied in 15 species of meso- and bathypelagic copepods from nine Calanoida families in the Arctic Ocean. During shipboard incubation, females of seven species released eggs freely into the water and females of three species produced membrane-bound egg sacs. One species, Aetideopsis rostrata, produced a mass of eggs looking like an egg sac, but the “sac” lacked a membrane and disintegrated within 2 h. Females of four additional species were encountered with membrane-bound egg sacs in the preserved samples. In most families, only one reproductive mode, either egg-carrying (the Euchaetidae, Augaptilidae), or broadcast spawning (the Heterorhabdidae, Spinocalanidae, Scolecitrichidae, Tharybidae, Bathypontiidae) was observed. In contrast, different genera of the Aetideidae family demonstrated different reproductive modes, with broadcast spawning predominant in the benthopelagic species, and both broadcast spawning and egg-brooding in the planktonic species. Clutch size and egg diameter varied widely between species in both broadcast spawners and egg-brooders. In broadcast spawners, the clutch size varied from 1 to 95 eggs female⁻¹, while the average egg diameter ranged from 152 to 440 μm. The clutch size for egg brooders varied between 3 and 82, while average egg diameter varied from 258 to 732 μm. Deep-water broadcast spawners produced much larger eggs compared to surface-dwelling broadcast spawning species. This larger egg size may result in a reduction, or elimination, of feeding during naupliar stages, thereby improving the survival potential of deep-water species.     

Plasticity in spawning visits of female damselfish, Stegastes nigricans : effect of distance to mates

Both sexes of the damselfish Stegastes nigricans hold individual territories in which they feed on filamentous algae. At dawn, females visit males' territories to spawn, and the males guard the eggs until hatching. We examined how females' spawning behavior varied according to the distances to their mates. Females usually mated with a single male per morning. The distance to the territory of a mate (0.7–12.8 m) did not affect a female's total spawning time per morning, but affected the number of her spawning visits with that male. Females made many repeated spawning visits when spawning with males at short distances, while they spawned the entire clutch in one visit when spawning with males at long distances. This plasticity in female behavior appears to be related to two costs during the spawning visits: (1) intrusions by other fish to feed on algae in the female's territory during her absence, which may cause the female to return repeatedly to her territory for defense, and (2) attacks on the female by other territorial fish, which increased with the distance to their mates' territories. To minimize the sum of both costs, females should change the number of spawning visits depending on the distance to the males' territories. Received: 30 September 1996 / Accepted after revision: 17 March 1997     

Reproduction and distribution of the invasive European fanworm Sabella spallanzanii (Polychaeta: Sabellidae) in Port Phillip Bay, Victoria, Australia

 The European fanworm Sabella spallanzanii (Gmelin, 1791) was recently introduced to Port Phillip Bay and is now a conspicuous component of most benthic communities. Reproduction of the worm was investigated in a population at Queenscliff over a 2 yr period (October 1995 to October 1997) using gonadal histology. The worms are dioecious (sex ratio 1:1, n=250), and attained sexual maturity at ∼50 mm body length. Reproductive periodicity followed a distinct annual cycle, and spawning proceeded through an extended autumn/winter period. Spawning was broadly synchronous between sexes, and coincided with falling seawater temperatures and shorter day-lengths. The females were highly fecund, and >50 000 eggs were probably shed from large females (>300 mm body length) during the annual spawning period. Breeding cycles of S. spallanzanii in Port Phillip Bay are ∼6 mo out of phase with endemic populations located at similar latitudes in the northern hemisphere. The spread of S. spallanzanii within Port Phillip Bay has been monitored by divers on an annual basis since 1994. The most recent dive survey (1998) indicates that S. spallanzanii has extended its range through out the entire 2000 km² embayment, and has invaded most subtidal habitats. Quantitative estimates of S. spallanzanii abundances were highest on pier pylons (12.5 individuals m⁻², 0.5 to 7 m depths). On sediments, estimates were highest at shallow sites (0.3 m⁻², 7 m depth), but numbers declined significantly with depth (0.1 m⁻², 17 to 22 m depth). Mean worm lengths and biomass were, by contrast, significantly higher at intermediate depths (12 to 17 m) than in shallower (7 m) or deeper (22 m) locations. S. spallanzanii demonstrates a clear preference for growth in sheltered, nutrient-enriched waters, so it may not spread from Port Phillip Bay into the adjacent oceanic waters of Bass Strait; however, in view of S. spallanzanii's current high abundance, fecundity and extended spawning periodicity, there is a high risk of future range expansions, mediated by shipping, into other temperate-water ports. Received: 17 November 1998 / Accepted: 6 January 2000     

Reproductive dynamics of endeavour prawns, Metapenaeus endeavouri and M. ensis, in Albatross Bay, Gulf of Carpentaria, Australia

 The spawning patterns of two penaeid prawns, Metapenaeus endeavouri (Schmitt) and M. ensis (De Haan), were examined from data collected at 45 stations between March 1986 and March 1992. An index of population fecundity based on the abundance, proportion and fecundity of sexually mature females was used as a measure of spawning output of the prawn stock. The population fecundity index for M. ensis was higher than that for M. endeavouri. The monthly population fecundity index for M. endeavouri varied markedly among years, while that for M. ensis was consistent among years. Spawning of M. endeavouri occurred year-round, while that of M. ensis was concentrated mainly in spring (September to November). For M. endeavouri, a minor spawning, derived from a relatively small number of summer spawners, occurred in the 20 to 30 m offshore waters in summer. In early summer (after May), the major spawning group consisted of large females from the winter-spawning cohort, and the spawning area shifted to depths of 30 to 60 m. In winter (July), the major spawning, derived from the winter-spawning cohort, occurred at depths of 20 to 40 m. For M. ensis, the major spawning, derived from the spring-spawning cohort, was observed in depths <50 m and was concentrated particularly in inshore waters (<20 m) in spring. In autumn, the spawning output was mainly from the autumn-spawning cohort, which comprised but a small number of individuals. In winter, the major spawning group again consisted of the large females from the spring-spawning cohort, and spawning increased in the oceanic waters (>50 m). These results suggest that mature female M. endeavouri and M. ensis move offshore (>40 m) by May and July, respectively, and return to shallow waters (<35 m) in July and November, respectively. The monthly reproduction patterns of both species in the “effective spawning” area showed that the major spawning season for M. endeavouri is August to October and that for M. ensis is September to December. Received: 19 February 1999 / Accepted: 18 June 2000     

Reproductive biology of the spiny lobster, <Emphasis Type="Italic">Panulirus Penicillatus</Emphasis>, in the southeastern coastal waters off Taiwan

The reproductive biology of spiny lobster, Panulirus penicillatus, was studied based on 2,068 lobsters, ranging from 34.28 to 131.60 mm carapace length (CL), sampled in Taitung coastal waters from September 2003 to December 2004. The overall sex ratio approximated 1:1 (χ² = 0.02, P > 0.05), but the monthly sex ratios in 2004 showed significant differences and males were predominant in sizes larger than 80 mm CL. Reproductive activity, assessed using histology, a gonadosomatic index and percentage of ovigerous females, indicated that the mature females could be found in every month and that the major spawning occurred from May to September. The presence of re-developing/re-ripe ovaries by month and size-specific spawning time suggest that larger mature females (>60 mm CL) spawn at least three times a year while smaller new mature females spawn at least once a year. For females, the estimated sizes at 50% physiological and functional maturity were (mean ± SE) 56.46 ± 0.56 mm CL and 66.63 ± 1.07 mm CL. The estimated sizes at functional maturity were between 72 and 74 mm CL for males. The number of eggs per spawning event (brood size, BS) was related to CL by the equation Y _BS = 2.4 × 10^-3CL^4.18 (r ² = 0.902, n = 12). Female lobsters with CL ranging from 60 to 80 mm made the greatest contributions to egg production because of their high brood size and active reproductive activity. A minimum legal size should be established for the fishery to protect egg production potential of lobster population in the southeastern coastal waters off Taiwan.     

Gonadal morphology and gametogenesis in the sea pen Pennatula aculeata (Anthozoa: Pennatulacea) from the Gulf of Maine

The ultrastructural features of gametogenesis are described in male and female colonies of the sea pen Pennatula aculeata. Specimens were collected for observation and fixation at 113 to 231 m depth in the Gulf of Maine, USA, in August 1993. The species is gonochoric, and all stages of gametogenesis are observed in both male and female colonies >45 mm in height. Gametogenesis shows several features that differ from sea anemones. The developing oocytes and sperm cysts are completely encompassed by gastrodermally derived follicle cells, and they are released from the mesenteries into the coelenteron before they are fully differentiated. Following maturation in the coelenteron, the eggs and intact sperm cysts are expelled through the mouths of the autozoids during spawning. The expulsion of sperm cysts suggests that they function as primitive spermatophores, perhaps as a way of reducing sperm dilution. Vitellogenesis results in the biosynthesis of lipid droplets which are the sole nutrient reserves in the egg. Heterosynthetic vitellogenesis is characterized by the importation of lipid precursors into the oocyte, and there is some indirect evidence that hypertrophic follicle cells play a role in production, transport, and/or mediation of these precursors. Spermatogenesis is similar to that of other anthozoans. The spermatozoon has a cone-shaped head, a posterior nuclear fossa, a ring of lipid-like bodies in the midpiece, a prominent cytoplasmic collar surrounding the proximal flagellum, and a single mitochondrion, but the posterior region of the sperm also contains previously undescribed concentric rings of cisternae resembling smooth endoplasmic reticulum. Received: 23 March 1998 / Accepted: 31 July 1998     

Longevity of subitaneous and diapause eggs of Centropages hamatus (Copepoda: Calanoida) from the northern Gulf of Mexico

Copepod resting eggs are abundant in the seabed of many bays and estuaries where they provide a potential source of recruits for growth of planktonic populations. In the northeastern Gulf of Mexico the copepod Centropages hamatus (Lillejeborg) occurs in the water column only during the late fall, winter and early spring. The species produces subitaneous and diapause eggs, and both egg types have been found in the seabed. We determined the longevity of these two egg types to ascertain their potential for contributing to the growth of the planktonic population and for sustaining a persistent egg bank. Eggs were collected from females and incubated in the laboratory under temperature and oxygen conditions chosen to simulate field conditions. The diapause eggs were also exposed to sulfide. The total hatching success of subitaneous eggs in two experiments declined from highs of 78 and 97% to zero after 60 and 90 d of exposure to anoxia. The total hatching success of diapause eggs that were exposed to anoxia for 90 d however was typically greater than 80%. Some diapause eggs hatched after being incubated under anoxia for 437 d. Diapause eggs survived longer at ambient field temperatures when incubated under anoxia (437 d) compared to normoxia (118 d). Exposure to sulfide did not result in greater mortality of diapause eggs compared to anoxia alone. Diapause eggs that were incubated at ambient field temperatures did not hatch when exposed to normoxia until the temperature dropped to <20 °C. The results of this study suggest that C.␣hamatus sustain a short-term reserve of subitaneous eggs in the seabed that provides recruits for the current year's population. The greater longevity of diapause eggs suggests that they sustain the seasonal reappearance of the species year after year in the northeastern Gulf of Mexico. However, the contribution of diapause eggs of C. hamatus from the Gulf of Mexico to a persistent egg bank is questionable since hatching ceased after 437 d. Received: 30 July 1997 / Accepted: 18 January 1998     

Reproductive biology in two species of deep-sea squids

Deep-sea squids, Moroteuthis ingens and Gonatus antarcticus, were collected in the slope waters off the Falkland Islands and their reproductive systems preserved and investigated onshore. Changes in oocyte length-frequencies at maturation and spawning, and their fecundity were studied. These squids, as well as many other species, are characterised by a synchronous oocyte growth and ovulation. Oviducts are not used for ripe egg accumulation and consequently the universal scale of Lipinski (1979) cannot be applied to assign female maturity. M. ingens spawns near the bottom; its fecundity is 168–297 thousand eggs. Maximum egg size is 1.8–2.7 mm. G. antarcticus spawns midwater; its fecundity is 10–25 thousand eggs. Egg size is 3.2–3.3 mm. In M. ingens spawning takes place in the austral autumn and winter, in G. antarcticus—in austral winter. Our data and the literature data show that the so-called “synchronous ovulation” probably occurs in all deepwater squids. This pattern is very rare among fish, but is quite common among benthic octopods that brood their egg masses.     

Effect of diet on reproductive characteristics of Ophryotrocha labronica (Polychaeta: Dorvilleidae)

Effects of food quality on fecundity (defined as the number of eggs deposited in the entire life of a female), egg size and egg energy content (measured by differential scanning calorimeter were studied in two populations of Ophryotrocha labronica (La Greca and Bacci): one taken from the natural environment immediately prior to the experiment, and the other from the same original environment, but bred for more than 5␣years in the laboratory. Results indicated that fecundity is mediated both by food quality and population origin. Specimens fed Tetramin generally spawned more frequently than those receiving the other two diets, while individuals bred in the laboratory since 1990 spawned more frequently than the wild population. Also the number of eggs per spawning and the total fecundity vary with the quality of the diet; Tetramin gives the highest fecundity values. The differences in egg size and energy content seem essentially related to the population origin, in fact, with all diets laboratory specimens produced larger eggs with higher energy contents than wild specimens. Received: 1 August 1997 / Accepted: 29 April 1998     

Maternal allocation of lipid classes and fatty acids with seasonal egg production in Atlantic cod (Gadus morhua) of wild origin

A suite of characteristics is often used to assess egg quality as these properties potentially play important roles in progeny survival and growth. Our objective was to assess egg characteristics including lipid biocomposition of an iteroparous, batch-spawning teleost of wild origin. Maternal allocation to egg number was generally dome-shaped (5 of 8 females) and egg size declined over the breeding season for eight breeding pairs of wild Atlantic cod (Gadus morhua) (n = 43 batches). Egg lipid composition ranged considerably among females and between egg batches within females (e.g., phospholipids 40–86 %; polar 47–87 % and neutral lipids 15–52 % of total lipids; polyunsaturated fatty acids 16–50 % of total fatty acids). Principal component analyses revealed significant inter-relationships among maternal traits, batch sequence and fecundity, and egg size and composition. Seasonal trends with regard to lipid deposition were variable; three females showed consistent declines in lipid parameters (μg egg⁻¹) with both batch number and egg diameter, one female showed consistent increase and the four remaining females showed no trend. The three females that exhibited seasonal declines in egg lipid content were characterized as having high fertilization success (>75 %). Our findings highlight the variability in lipid allocation to eggs of batch spawners of wild origin and characterize the composition of endogenous reserves available during embryogenesis and yolk sac larval stages.     

Population biology of Dyopedos monacanthus (Crustacea: Amphipoda) on estuarine soft-bottoms: importance of extended parental care and pelagic movements

The suspension-feeding amphipod Dyopedos monacanthus (Metzger, 1875) is a common epibenthic amphipod that lives on self-constructed “mud whips” (built from filamentous algae, detritus and sediment particles) in estuaries of the northern North Atlantic Ocean. The population biology of D. monacanthus at a shallow subtidal site in the Damariscotta River Estuary (Maine, USA) was examined between July 1995 and July 1997. The resident population at the study site was dominated by adult females during most months of the year. High percentages of subadults were found in late summer/early fall. Often, between 10 and 20% of the adult females were paired with males, and the percentage of ovigerous females varied between 40 and 100%, indicating continuous reproduction. The percentage of parental females varied between 40 and 80% during most months, but dropped to levels below 20% during summer/early fall. The average size of amphipods on their own mud whips was ∼4 mm during the summer/early fall, after which it increased continuously to >7.0 mm in March or April, and then dropped again. In March and April, the average number of eggs and juveniles female⁻¹ was ∼100 eggs and 55 juveniles, while during the summer/early fall the average number of eggs female⁻¹ was <20 and that of juveniles female⁻¹ was <10. Many juveniles grew to large sizes (>1.4 mm) on their mothers' whips in winter/early spring but not in the summer/fall. The average number of amphipods at the study site was low in late summer/early fall (<50 individuals m⁻²), increased steadily during the winter, and reached peak densities of >3000 individuals m⁻² in April 1996 (>1600 individuals m⁻² in May 1997), after which densities decreased again. The decrease of the D.␣monacanthus population at the study site coincided with a strong increase of amphipods found pelagic in the water column. This behavioural shift occurred when temperatures increased and benthic predators became more abundant and active on shallow soft-bottoms, suggesting that D. monacanthus at the study site is strongly affected by predation. The effects are direct (by predation on amphipods) and indirect (by reducing duration of extended parental care and enhancing pelagic movements). Both extended parental care and pelagic movements are important behavioural traits of D.␣monacanthus (and other marine amphipods), and significantly affect its population dynamics. Received: 18 January 1998 / Accepted: 27 May 1998     

Population dynamics and production of the planktonic copepods in a eutrophic inlet of the Inland Sea of Japan. IV. Pseudodiaptomus marinus, the egg-carrying calanoid

Population dynamics and production of the egg-carrying calanoid copepod Pseudodiaptomus marinus were studied for a year in Fukuyama Harbor, a eutrophic inlet of the Inland Sea of Japan. This species was perennial, with a large numerical peak in June and small peaks in September/October and November/December. During the study period, at least 11 generations could be detected. For each generation, the stage-specific survival from egg to Copepodite Stage (C) V was determined; it was very high during early life stages (egg to NIII), and gradually decreased beyond. On average, 94% of eggs recruited into NIII, which is strongly contrasted with very high (>ca. 90%) mortality during the corresponding stages for free-spawning copepods, i.e. Acartia omorii, Centropages abdominalis and Paracalans sp. This demonstrates that the egg-carrying strategy has a great advantage to reduce mortality in egg stage. The biomass of this species showed marked seasonal variations largely in parallel with numerical abundance. The instantaneous somatic growth rate increased linearly with temperature. The population production rate was estimated as the sum of somatic growth of larval stages and egg production of adult females; the annual integration was 51.0 mg C m⁻³ yr⁻¹ or 0.38 g C m⁻² yr⁻¹. Received: 11 November 1996 / Accepted: 7 December 1996     

Vertical distribution, life cycle, and developmental characteristics of the mesopelagic calanoid copepod Gaidius variabilis (Aetideidae) in the Oyashio region, western North Pacific Ocean

Vertical distribution, life cycle, and developmental characteristics of the mesopelagic copepod Gaidius variabilis Brodsky in the Oyashio region were investigated by combining analyses of field copepodite populations with laboratory-rearing data of egg hatching and naupliar development. Field samplings from five discrete depths between the surface and ≤2000 m were made approximately every month for 1 year. Most populations of G. variabilis occurred between 600 and 1000 m depth. A modest degree of reversed diel vertical migration behavior and some stage-specific depth-distribution patterns were noted. All copepodite stages were observed throughout the year, suggesting a year-round spawning of G. variabilis. From a prominent abundance peak of Copepodite Stage 1 (C1) seen in June to August, together with development times of eggs and nauplii obtained in laboratory-rearing experiments, the major spawning season was extrapolated to be April to June, the phytoplankton bloom season. Tracing the peak abundance of each copepodite stage (distinguishing males and females for C4 to C6), the generation times of males and females were deduced as 2 and 1 year, respectively. All between-stage increments in terms of wet-, dry-, and ash-free dry weights were greatest in C3/C4, and least in C5/C6 for both males and females. The increments in C3/C4 and C4/C5 were greater for males than for females, reflecting a longer stage duration of the males. These weights did not increase in C5/C6 males, possibly because feeding ceased in C6 males. These results for G. variabilis are compared with those for some mesopelagic copepods previously reported from other regions. Received: 25 October 1999 / Accepted: 20 March 2000     

Changes in metabolic substrates during early development in anchoveta Engraulis ringens (Jenyns 1842) in the Humboldt Current

We assessed the ontogenetic changes in protein content and free amino acids (FAA) in eggs and early larvae of Engraulis ringens (anchoveta) off central Chile on different dates during the spawning season. On all sampling dates, a reduction in embryonic yolk-sac volume, proteins and FAA concentrations occurred during development. Protein electrophoresis (SDS–PAGE) of eggs and larvae showed at least 22 protein bands: 11 were consumed early and not detected after hatching. The proportion of essential FAA (EFAA) was higher than the proportion of non-essential FAA (NEFAA) in early eggs and in 7 day-old larvae (82.5-73% EFAA respectively). During egg development, the FAA pool was dominated by leucine, alanine and lysine, three amino acids contributing 35–44% of the total FAA in eggs. During larval development, histidine was the most abundant FAA. In July, total FAA constituted 13–18% of the egg dry weight. A similar proportion (45–51%) occurred in July between protein plus FAA and total lipids. The differences in egg size during the spawning season along with variability in batch composition suggests that the female spawning condition is a major factor determining egg quality and early offspring success.