Reproductive biology of the common antarctic crinoid Promachocrinus kerguelensis (Echinodermata: Crinoidea)

Histological examinations of the gonads of Promachocrinus kerguelensis Carpenter, 1888 collected in November 1984 from McMurdo Sound, Antarctica, revealed ripe ovaries containing three distinct size classes of oocytes: 20 to 90, 91 to 150 and 151 to 180 m diameter as well as ova. Testes were ripe, and filled with mature sperms. There was marked inter-pinnule and inter-individual synchrony of gametogenesis and spawning is probably synchronous, at least within the sub-population sampled. The number of eggs per pinnule is allometrically related to body size. Body weight and arm length are positively correlated with total egg number. Annual fecundity was estimated to be 29 000 eggs (full-grown oocytes and ova) for a crinoid weighing 53.6 g wet weight with an arm length of 16.3 cm. The eggs float. Female genital pinnules contain more lipid and energy than male pinnules. Reproductive output and effort (23.4 to 65.2 kJ and 26 to 38%, respectively) increase with body size and are similar to values reported for other echinoderms with lecithotrophic larvae. The greatest overall energetic contribution is represented by the arms and genital pinnules. Spawning in P. kerguelensis probably occurs in November and December within McMurdo Sound. If development is slow, as in all other antarctic echinoderms studied to date, then settlement most likely occurs 2 to 3 mo later, after the short summer period of high phytoplankton productivity. High fecundity, a pelagic mode of reproduction, and the high probability of a slow rate of development may contribute to the wide geographic distribution of this common antarctic crinoid.     

Reproduction and condition of female Octopus mimus (Mollusca: Cephalopoda)

The relationship between reproduction and condition was studied in a 15 mo sample of 919 maturing, mature and post-spawning female Octopus mimus (388 to 3714 g) caught in Iquique (North Chile). O. mimus is a semelparous species, with reproduction taking place all year round. Investment in reproductive tissues was, on average, 9.9% of mature female body weight, independent of season. However, somatic growth during maturation was dependent on season and varied between 26 and 63% of the whole-body growth in weight. The condition of females did not vary markedly until spawning, although seasonal variations were apparent, winter being the most unfavourable. Condition deteriorated dramatically after spawning, during parental care of the eggs. During this period, somatic-tissue depletion, mainly from the muscles, was >25% of the total body weight of mature females. The fecundity of O. mimus was probably limited by the costs associated with parental care of the eggs.     

Social inhibition of maturation in females of the temperate wrasse Pseudolabrus celidotus and a comparison with the blennioid Tripterygion varium

In two temperate reef fish, Pseudolabrus celidotus (Bloch and Schneider) and Tripterygion varium (Bloch and Schneider) studied near Leigh, New Zealand, most spawning activity was concentrated during the first 2 months of spawning seasons which lasted about 5 months. In P. celidotus, maturation and spawning of first-year females (0+) was delayed with respect to older (1+yr) females. Within the 0+ age class, the minimum size of maturation declined as the spawning season proceeded. However, in all females of T. varium, ripening and spawning took place over the same time ranges. Observations on the seasonal patterns of female aggression suggested that larger female P. celidotus were socially inhibiting the maturation of small females. The level of interaction was high compared to that in T. varium, and reached a peak over the onset of the spawning season. This hypothesis was tested by the removal of larger (1+yr) female P. celidotus from a field population. The remaining 0+ females increased in ovary weight and matured earlier than those in an undisturbed area. We argue from this experiment that it is of advantage for P. celidotus females to spawn early in the season and of advantage to inhibit other females from doing so. Two possible reasons are suggested, the first applying to sexchanging species such as P. celidotus, the second to temperate-water fishes in general.     

Maturity classification and group maturity of the red sea bream Pagrus major

The criteria of ovarian maturity during the spawning season and maturity rates according to age are described for the female red sea bream Pagrus major collected from the Chikuzenkai between April and June, 1984. The degree of maturity of the ovary is categorized into four stages based on the maturation stage of the most advanced oocytes in the ovary, rate of atresia and occurrence of post-ovulatory follicles. The ovarian maturity rates in fish aged 2, 3, 4 and 5 years were shown to be 5.8, 46.2, 75.0 and 97.3%, respectively, assuming that yolked ovaries with low levels of atresia (<50% yolked oocytes affected) attain maturity. The fork length at which 50% of the fish attained maturity ranged from 280 to 300 mm.     

Spawning pattern and type of fecundity in relation to ovarian allometry in the round herring <Emphasis Type="Italic">Etrumeus teres</Emphasis>

Spawning pattern (assessed by seasonal changes in ovarian developmental stages) and type of fecundity (assessed by analysis of oocyte-size frequency distributions) of the round herring Etrumeus teres were studied in relation to ovarian growth and seasonal changes in the gonadosomatic (GSI), hepatosomatic (HSI) and liposomatic (LSI) index as well as the somatic condition of spawners (CS) in a spawning ground of southern Japan. Except for summer, mature and recently spawned ovaries occurred all year round. Oogonia and primary oocytes were present in all ovaries, and cortical alveoli stage (CA) oocytes occurred in all mature, hydrated and partially spent (PS) females (PS: females containing post-ovulatory follicles). Before hydration, a clutch of larger yolked oocytes, undergoing synchronous growth (range 0.7–1.1 mm), was present in mature ovaries which was completely separated from a more heterogeneous clutch of oogonia, primary and secondary oocytes (<0.150 mm) and oocytes in the CA stage (range 0.15–0.60 mm). As vitellogenesis progressed, the yolked clutch increased in size but the CA oocytes remained arrested. The latter entered into the secondary growth phase when hydration started in the advanced batch. Ovarian growth was isometric in all developmental stages, validating the use of GSI, which showed a consistent monthly evolution among years. Spawning stopped in summer (July and August) and peaked in winter and spring. HSI correlated positively with GSI on both a monthly mean basis (r = 0.76) and individual fish basis (liver weight explained 67–83% of the variability in ovary weight when females were grouped into 1-unit GSI intervals) suggesting a significant role of liver in vitellogenesis. LSI and CS also showed marked seasonal changes peaking from summer to middle autumn. Overall results suggest that E. teres is a multiple spawner with a group-synchronous ovarian development and indeterminate annual fecundity, with the three processes linked to an isometric growth of the ovary. We propose that such a reproductive pattern is an adaptation to produce batches of large pelagic eggs through a protracted spawning season.     

Contribution of moulting and eggs to secondary production in Nyctiphanes australis (Crustacea: Euphausiacea)

This study was designed to assess the contribution of moulting and eggs to production of the euphausiid Nyctiphanes australis G. O. Sars. For this purpose, live specimens were collected from Storm Bay, south-eastern Tasmania, between August and December 1981, while preserved samples, collected between February 1980 and February 1981, were also examined. The intermoult period of N. australis increased exponentially with increase in body length and weight. Larvae moulted approximately every 2.5 to 3 d and adults every 4 to 5 d at 15°C. A decrease in temperature from 15° to 10°C resulted in the intermoult period almost doubling. The mean weight of exuviae produced represented nearly 6% of the body dry weight of the individual. Continuous maturation of ova was observed, with an individual female capable of releasing a total of 1 100 eggs in a lifetime. These are deposited as a series of batches into a pair of external ovisacs every 30 d. The size of the batch of eggs was dependent on the size of the female. Moulting of euphausiids forms a significant contribution of organic matter to detrital food webs. A value of 42.01 mg m^-3 yr^-1 was obtained for the production of exuviae. Egg production was calculated to be between 1.41 and 4.22 mg m^-3 yr^-1. Production and mean annual biomass of N. australis was previously calculated as 78.29 mg m^-3 yr^-1 and 5.39 mg m^-3, respectively. Thus, the total production integrated for the whole of Storm Bay was 125 mg m^-3 yr^-1 or 2 309 tonnes dry wt yr^-1, representing an overall P:B ratio of 23.1.     

Effect of intraspecific density on life history traits and population growth rate of Neanthes arenaceodentata (Polychaeta: Nereidae) in the laboratory

The effects of intraspecific density on life history traits and population dynamics of the nereid polychaete Neanthes arenaceodentata Moore were assessed in a laboratory experiment. Survival, growth, and fecundity were measured for one generation of worms at densities of 40, 80 and 160 worms per 840 cm² (1x, 2x, and 4x treatments, respectively). Density did not affect size (prior to pairing), percentage of worms paired, time to pairing, or size of mature paired males. Density did have a significant negative effect on survival, size of mature paired females, time to spawning, percentage of females that reproduced, and number of eggs per reproducing female. As density increased, mean survival was 90.0, 80.8 and 74.0%; mean size of mature females was 52.2, 49.2 and 48.1 segments; mean time to spawning was 100.6, 102.4 and 109.4 d; and mean fecundity was 881, 622, and 598 eggs per female, for 1x, 2x and 4x treatments, respectively. Increased density reduced the potential population growth rate, ; for a given rate of larval survivorship, was lower in the 2x and 4x treatments than the 1x treatment. Analysis of sensitivity of to changes in survivorship indicated that population growth rate at the highest density was sensitive to both changes in larval survivorship and the probability of producing a successful brood, although at low densities, was sensitive only to changes in larval survivorship. We attribute these density effects to aggressive bahavioral interactions between the worms, primarily the adults. This experiment identifies key life history traits that could be measured in future experiments to test population level responses of N. arenaceodentata populations to pollutants, both in the laboratory and in the field.Contribution No. 820 of the U.S. Environmental Protection Agency, Environmental Research Laboratory and Contribution No. 189 of the Marine Sciences Institute, University of Connecticut     

Reef coral reproduction in the eastern Pacific: Costa Rica,Panamá, and the Galápagos Islands (Ecuador). VI. Agariciidae, <Emphasis Type="Italic">Pavona clavus</Emphasis>

The reproductive ecology of the zooxanthellate reef coral Pavona clavus was investigated at several sites in Costa Rica, Panamá, and the Galápagos Islands (Ecuador) over the period 1985–2009. Pavona clavus displayed stable gonochorism as only five hermaphrodites were found in 590 samples. At four of five locations, sex ratios were skewed toward female dominance; however, at Saboga Island (Panamá) male colonies predominated. In Panamá, sexual maturity was observed in an estimated eight-year-old female colony, and several colonies of 10–20 years of age demonstrated gametogenesis. Sexual activity was observed at all study sites, but gamete development occurred in only 14–31% of colonies sampled sporadically. Seasonality of gametogenic activity occurred predominantly during the warm/wet season, June to August, at mainland sites (Caño Island, Costa Rica, and Gulfs of Chiriquí and Panamá, Panamá). This pattern was repeated in the Galápagos Islands, but mainly from March to May when seasonally high sea temperatures and rainfall prevailed there. Histological sampling and field observations indicated that spawning was centered around the full moon, most frequently on lunar day 17, and near sunset (1,800 h). Mean fecundity (mature ova cm^?2 live tissue) estimates were significantly different for two sites and ranged from ~1,780 (Saboga Island, Gulf of Panamá, seasonally upwelling) to ~4,280 (Uva Is, Gulf of Chiriquí, non-upwelling). Assuming three annual spawning events colony^?1 (August, September, October), extrapolation of minimum and maximum fecundities yield 5,340 and 12,840 ova cm^?2 year^?1. Seasonal, lunar, and diel spawning patterns in nine zooxanthellate species at Uva Island indicate asynchronous coral community spawning.     

Does the spawning of Calanus finmarchicus in high latitudes follow a reproducible pattern?

In the spring of 1989, an experimental study of the spawning behaviour of Calanus finmarchicus was carried out in Malangen, northern Norway. Here, a single cohort of females reproduce from mid-March to May, approximately coinciding with the wax and wane of the spring phytoplankton bloom. An evaluation of population characteristics such as the proportion of adults, sex ratio, as well as gonad maturation and daily productivity of the females clearly reveals three phases within the population's reproductive period. In between incline and decline, the highest spawning rates (on average >20 eggs female^-1 d^-1, equivalent to 5.7% body C d^-1) occur after the males have disappeared from the population and almost all females have mature gonads. During this period, the ratio of adults to copepodid Stage Vs changes from dominance of adults to that of CVs. Although first egg production was observed prior to the phytoplankton increase, it is suggested that the onset of the phytoplankton spring bloom in the first few days of April enhances the final maturation of ovaries in the females and therefore triggers the onset of the main spawning period. The clutch sizes (max. 95 eggs clutch^-1) vary with the age of the females, while the spawning frequencies depend on the available food quantities. The overlap of an estimated minimal 4 wk spawning period for the individuals leads to a main reproductive phase for the population of ca. 3 wk, during which time mean clutch sizes and spawning frequencies are maximal (highest average clutch size: 70 eggs female^-1 clutch^-1, 100 to 60% of the females spawning). This period ends before the end of the phytoplankton bloom. Calculated by stepwise interpolation and summation of the mean daily egg production in the population, an average female produced ca. 600 eggs during the spring bloom in Malangen 1989. We suggest that reproduction and population development of C. finmarchicus in spring follows a reproducible pattern for a given temperature regime and non-limiting food conditions. In the case of clearly identifiable cohorts, it seems possible to trace the state of reproduction by evaluating population parameters.     

Reproduction and reproductive variability in the coral Pocillopora verrucosa from the Republic of Maldives

The sexuality, reproductive mode, and timing of reproduction of Pocillopora verrucosa from the Republic of Maldives, Indian Ocean, were assessed using serial histological sections. These showed that P. verrucosa is an annual simultaneous hermaphrodite, with gonads arranged in two opposing arcs of alternating testes and ovaries, six gonads in each arc. No planulae were observed in any dissection or histological analysis carried out, therefore making brooding unlikely. Broadcast spawning is inferred from the disappearance of mature gametes from samples collected between late March and April 1991. Mean oocyte size at spawning was 53.5 m and mean potential fecundity was 7300 ooctyes cm^-2yr^-1. The reproductive pattern of P. verrucosa in the Maldives is compared to that of the same species in different locations, the short breeding season in March to April occurring earlier in the Maldives than in Red Sea populations. The maximum mean oocyte diameter found in Maldivian specimens was much smaller than elsewhere. The year-to-year variation in numbers of oocytes/polyp within single colonies, and the variation between colonies within the population was significant (p<0.01). Therefore, it is possible that fecundity is not a good variable to use when monitoring stress on coral reefs unless larger numbers of estimates can be routinely made. Some colonies contained immature oocytes at spawning that were not released and continued to grow to approximately twice the size of the spawned oocytes that were presumed to be mature. These unspawned oocytes were oosorbing, and were characterised by the presence of zooxanthellae and large numbers of vacuoles in their cytoplasm. This gradual expansion and oosorption of unspawned oocytes has not been documented previously for corals.     

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     

Maturation,fecundity, and spawning strategy of the thornback ray, <Emphasis Type="Italic">Raja clavata</Emphasis>: do reproductive characteristics vary regionally?

In Portuguese waters, thornback ray Raja clavata spawns mainly between May and January, although females and males in spawning condition are found throughout the year. The maturation process can be divided into three main phases by using information on gonad weight, oviducal gland and uterus width in females and on gonad weight, clasper length and sperm duct width in males. Females attain length-at-first-maturity at 784 mm and males at 676 mm, at ages of 7.5 and 5.8 years, respectively. In females larger than length-at-first-maturity, a resting stage was identified characterized by low gonadosomatic index and well-developed oviducal glands and uteri. These results along with the low proportion of adult females that are effectively reproducing per month demonstrated that the thornback ray cannot be considered a continuous spawner as described in other studies. Fecundity was determinate with about 35 eggs released per batch. During the spawning season, a total of four batch episodes occur indicating that the total fecundity was approximately 136 eggs per female. Regional differences may exist in the reproductive strategy of the species, namely on the duration of the spawning season, length-at-first-maturity, and fecundity, which can be related to a more intense fishing pressure in northern European waters.     

Mating behaviour and reproductive cycle of Archaster typicus (Echinodermata: Asteroidea)

This study examines the reproductive cycle, the mechanism of male-on-female pairing behavior and the spawning behavior of Archaster typicus Müller et Troschel. Field studies were conducted in the intertidal zone of the sand beaches at Penghu, Taiwan (23 32N; 119 33E) at ebb-tide in 1984 and 1985. The pairing behavior of A. typicus is a reproductive behavior which leads to simultaneous spawning, increasing the probability of fertilization. As the breeding season approaches, sea stars, especially males, display increased mobility. Because only males tend to mount another individual and because males can detect the sex of another individual by contact with the side of their arms, a male-on-female pair is formed when a male encounters a female. Eighty-five percent of the sea stars observed were paired during the pairing season. Spawning by a paired female is closely followed by spawning of its paired male; male spawning, however, does not induce spawning in its paired female. During spawning, the male turns slightly so that its arms overlap the arched arms of the female. The gonad volume of males is much less than that of females. This may result from the high efficiency of fertilization in this species, which does not require a large amount of gametes to be released, or from the higher energetic demands made on the males.     

Egg production by the subtropical copepod Undinula vulgaris

Egg production by Undinula vulgaris, collected off Kaneohe Bay, Hawaii, was measured under field and laboratory conditions on 16 occasions from June to December 1991. In situ rates ranged from 0 to 15.7 eggs female^-1 d^-1, with a mean of 6.4 eggs female^-1 d^-1, 2.1% female body carbon d^-1. Maximum in situ production was 53 eggs female^-1 d^-1, 17.2% C d^-1. Average egg production (Y, eggs female^-1 d^-1) was related to the concentration of particulate carbon (X, g C l^-1) by the Ivlev function, Y=13.9[1-e^{-0.0097 (x-10)}], with R ²=0.96. Individuals with the same feeding history produced more eggs at lower temperatures in the laboratory. Egg production was not significantly correlated with dry weight, and no noticeable temporal trend was found. Despite the elevated habitat temperatures (26 to 27°C) of this subtropical copepod, maximum fecundity of U. vulgaris was comparable to, but average rates were lower than, egg production rates of similarly-sized, temperate and borcal species of the genus Calanus. Our results caution against broad extrapolations of the temperature-growth relationship for temperate coastal copepods to species from poorly studied, oligotrophic regions of the oceans.Contribution No. 3257 from the School of Ocean and Earth Science and Technology, University of Hawaii at Manoa, Honolulu, Hawaii 96822, USA     

Developmental types of shallow-water asteroids of McMurdo Sound,Antarctica

The mode of development was ascertained for 14 of the 16 species of sea stars known to occur in shallow waters of McMurdo Sound, Antarctica (77°51S; 166°40E). The species were collected between September 1984 and December 1985. Females of three species,Odontaster validus, O. meridionalis andPorania antarctica, spawn small to moderate eggs (0.17 to 0.55 mm), have a high fecundity, and produce feeding larvae. Females of an undescribedPorania species spawn a few eggs (150 to 310) that are 0.55 mm in diameter and develop into demersal non-feeding larvae. Females ofDiplasterias brucei andNotasterias armata produce a few (<300) large eggs (2.8 to 3.5 mm) and brood their young. Females of the remaining eight species have moderate fecundity and produce pelagic non-feeding larvae, as determined from egg type (buoyant, 0.54 to 1.28 mm diam) and direct observations of spawning and development. The high incidence (11 out of 14 species; 79%) of non-feeding development is consistent with predictions that environmental conditions in high-latitude regions are unfavorable for planktotrophic development. Nonetheless, most of the species surveyed (11 out of 14) had pelagic larvae, which contradicts inferences of unusual selection for benthic development in the Antarctic.     

Comparison of larval development and growth of the sea cucumberActinopyga echinites: Ovary-induced ova and DTT-induced ova

AdultActinopyga echinites (Jaeger) were collected from northern Taiwan in September 1989. Oocytes were induced to mature by bathing them in ovary juice (ovary-induced ova) or in 10^–2 M dithiothreitol (DTT-induced ova). The percentage of germinal vesicle breakdown (GVBD) increased from 0.4 to 6.4% in the former treatment and to 84% in the latter. After artificial fertilization, the embryos were cultured in seawater (35 S) at 25 to 28°C. Larvae were fed with the algaIsochysis aff.galbana at 10^4–5 cells/ml. Larvae from ovary-induced ova developed faster (18 d to the doliolaria stage) and grew to a larger size (1.13 mm length) than those from DTT-induced ova (20 d to the doliolaria stage and 0.62 mm in length). On the twelfth day, larvae from ovary-induced ova bear lipid spheres. The number of spheres is positively correlated with larval size. Lipid spheres may provide nutrient reserves for larvae during metamorphosis.     

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 parameters of female Hawaiian damselfish<Emphasis Type="Italic"> Dascyllus albisella</Emphasis> with comparison to other tropical and subtropical damselfishes

The reproductive ecology of female Hawaiian damselfish Dascyllus albisella (Gill) was studied at Kaneohe Bay, Oahu, Hawaii (21°27'N; 157°47'W) during two peak reproductive seasons (June–September 1997, July–August 1998). In both years, spawning occurred cyclically every 5–7 days, with all spawning in the study area in each cycle concluding within 2–3 days. The study period encompassed 16 spawning cycles in 1997 and 10 cycles in 1998. Some females spawned in almost all cycles, whereas others spawned in as few as one. Mean spawning interval of individual females ranged from 6.4 to 11.7 days in 1997 and from 5.5 to 29.0 days in 1998. Batch fecundity increased exponentially with body length, and ranged from 23,100 to 52,800. Spawning duration increased linearly with body length and ranged from 20 to 286 min. On a single day, large females spawned earlier than small females. Females mated with several different males over a season, but typically mated with only one male on a single spawning day. Female spawning frequency and interval, spawning duration, batch fecundity, as well as diel timing of spawning in D. albisella were comparable to those reported for other large, planktivorous, non-territorial damselfishes.     

Life history of the sandy-beach amphipod Neohaustorius schmitzi (Crustacea: Haustoriidae)

Neohaustorius schmitzi Bousfield, the dominant macroscopic invertebrate on sandy beaches in North Carolina, USA, is strictly intertidal, and distributed from mean low tide to high tide, with an average density of 790/m². Location of greatest densities within the intertidal zone is dependent upon season and sex. An aggregated distribution is shown. N. schmitzi has 2 generations/year: a winter generation which lives about 8 months, and a summer generation which lives about 4 months. The reproductive season extends from February through October, with spring and summer peaks. Each female appears to produce only 1 brood of young. The number of young ranges from 2 to 14, depending upon the length and generation of the female.     

Life history of the golden ring cowry Cypraea annulus (Mollusca: gastropoda) on Okinawa Island,Japan

Two mark-recapture studies, regular population censuses, field observations, and laboratory culture were used to study the life history of the tropical marine gastropod Cypraea annulus Linnaeus, 1758 from Cape Maeda, Okinawa Island, from April 1984 to March 1986. In the field, the average rate of increase in shell length of marked juvenile snails was 1.0±0.3 mm wk^-1 (N=13), with a maximum of 1.5 mm wk^-1 (initial shell length 10.6 to 17.2 mm). In the laboratory, the maximum growth rate of juveniles was 3.0 mm wk^-1 with food ad libitum. Snails with primordial teeth on the shells grew at a rate of 0.1 to 0.5 mm wk^-1 for about 2 wk. The adults continued to grow at a similar rate for an additional 3 to 6 wk, and ceased detectable growth when some females started spawning egg masses. In the littoral zone on Okinawa Island, snails reproduced throughout the year. Egg masses brooded by three females (shell length 17.4 to 21.3 mm) in the field contained 90 000 to 133 000 ova. In the laboratory, brooding periods of three females lasted 6,8, and 9 d. The estimated average frequency of spawning was 5 egg masses female^-1 yr^-1. The mean shell length of adult females (20.3 mm) was significantly larger than that of adult males (19.6 mm). The life-history strategy of C. annulus is characterized by rapid growth, high fecundity, and repeated spawning throughout the year.