Observations on the life history of the blue-ringed octopus Hapalochlaena maculosa

The life-history of the blue-ringed octopus Hapalochlaena maculosa (Hoyle) was observed in laboratory aquaria. Eggs from a brooding female were bred through to the next generation. The life cycle lasts approximately 7 months–4 months from hatching to maturity, 1 month from copulation to egg-laying, and an estimated 2 months for embryonic development. This venomous octopus has several unique and interesting habits. Eggs are not attached to a substratum but are carried by the mother throughout their embryonic life. She assumes a brooding posture for the greater part of the time and cradles the eggs between her raised skirt and body. However, she can move froely with her clutch of eggs, conveying them individually or in clusters along her arms from one sucker to the next as occasion demands. Embryos reverse position within their capsules 1 week prior to hatching, and hatch at the distal end of the capsule, mantle-first. Development is direct; there is no planktonic stage. The young immediately assume a benthic habit and, within a few hours, consume the remnant of the yolk-sac which they carry with them from the egg. Juveniles begin to feed on pieces of crab within a week of hatching, and to kill and eat live crabs within a month. They pierce the carapace of their prey at the abdominal articulation and with the aid, apparently, of venom from their posterior salivary glands suck out the partially pre-digested tissues. The ink gland is relatively small in size and is mainly used between the second and the fourth week of juvenile life. Chromatophores begin to function during embryonic life. After hatching there is a progressive change in coloration, and the iridescent blue rings which characterize the species appear at the age of about 6 weeks. Mating is a more active process than has been recorded for other octopuses. The male mounts the female and clasps her securely. After a short struggle the female becomes quite passive, almost inert. Copulation lasts about one hour. The unusual life-history of H. maculosa suggests that it is a highly evolved octopus species. Its direct development, simple diet, and rapid rate of growth make the animal relatively easy to cultivate for experimental or pharmacological purposes.     

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     

Laboratory observations on spawning,fecundity and early development of a mesopelagic ostracod,Conchoecia pseudodiscophora,from the Japan Sea

The fate of the polycyclic aromatic hydrocarbon [³H]benzo[a]pyrene (BaP) was examined in two species of scleractinian corals, Favia fragum (Esper) and Montastrea annularis (Ellis and Solander), which were collected in the patch reefs surrounding Alina's Reef (25°23.25N; 80°09.8W) in Biscayne National Park, Florida, USA, in July, 1990. Corals were exposed to initial concentrations of 5 g/l in a simple static system for 25 h. BaP uptake was estimated from the disappearance of BaP from the water. Uptake rates were 6.5±0.7 and 10.8±0.2 g BaP cm^-2h^-1 for F. fragum and M. annularis, respectively, at initial BaP concentrations and were directly proportional to the concentration of BaP in the water. The separation of zooxanthellae from coral tissue revealed that zooxanthellae can accumulate up to 53 and 64% of the total BaP-derived radioactivity present in F. fragum and M. annularis, respectively. Both corals metabolized BaP slowly, as most of the accumulated radioactivity was present as the unmetabolized chemical. However, aqueous and organic-soluble metabolites were found in both the animal and zooxanthellae fractions. Analysis by high-performance liquid chromatography (HPLC) revealed that both species of corals metabolized BaP to various tetrols, triols, dihydrodiols, quinones and phenols, although the pattern of metabolites differed between species. Zooxanthellae contained some of the same Phase I metabolites found in the animal tissue; however, tetrols and triols were absent in extracts from the zooxanthellae. The elimination of BAP from corals was also slow; approximately 38 and 65% of the accumulated radioactivity was still present in F. fragum and M. annularis, respectively, 144 h following the transfer of exposed corals to an uncontaminated flow-through seawater system.     

Chemical defense in pelagic octopus paralarvae: Tetrodotoxin alone does not protect individual paralarvae of the greater blue-ringed octopus (Hapalochlaena lunulata) from common reef predators

Some pelagic marine larvae possess anti-predator chemical defenses. Occasionally, toxic adults imbue their young with their own defensive cocktails. We examined paralarvae of the greater blue-ringed octopus (Hapalochlaena lunulata) for the deadly neurotoxin tetrodotoxin (TTX), and if present, whether TTX conferred protection to individual paralarvae. Paralarvae of H. lunulata possessed 150 ± 17 ng TTX each. These paralarvae appeared distasteful to a variety of fish and stomatopod predators, yet food items spiked with 200 ng TTX were readily consumed by predators. We conclude that TTX alone does not confer individual protection to paralarvae of H. lunulata, and that they possess an alternative defense. In larger doses, tetrodotoxin is a deterrent to the predatory stomatopod Haptosquilla trispinosa (mean dose = 3.97 μg/g). This corresponds to 12–13 paralarvae per predator based on the TTX levels of the clutch we examined. Thus, the basic assumption that individual paralarvae of H. lunulata are defended by TTX alone was disproved. Instead, functionality of TTX levels in paralarvae may arise through alternative selective pathways, such as deterrence to parasites, through kin selection, or against predator species not tested here.     

Laboratory observations on spawning,brood size and egg hatchability of the Antarctic krill Euphausia superba from Prydz Bay,Antarctica

Forty-three female Euphausia superba were collected from three sites in Prydz Bay, Antarctica in January 1985. Most gravid females took 3 to 14 d to spawn after capture and some less mature females released eggs after 21 d. Most females spawned only once; overall mean brood size was 2 809 eggs and mean hatchability for eggs from the premoult females was 69.7%. For seven females spawning a second time, overall mean brood size was 597 eggs. Mean hatchability for second-spawn eggs from five premoult females was 67.1%. Possible origins of the second-spawned eggs were small numbers of matured eggs left in the ovaries after the first spawning or eggs developed from those of peripheral nucleolus and vitellogenic stages. Throughout the 8 wk of observations, there was no apparent development of eggs in females which had released all their eggs at one spawning.     

Some observations on spawning and fecundity in the luminescent fish Photoblepharon palpebratus

Spawning pairs of the fish Photoblepharon palpebratus (Boddaert), a bioluminescent shallow-water species from the Banda Islands, were observed in captivity for the first time. Both female individuals of the two pairs studied were slightly larger than the males, and possessed rounder tail-fin edges, criteria which can be used for sexing. Laboratory and field observations indicate that during several dark nights in April and May one female could produce a total of up to 1,000 sticky, transparent, 1.2-mm diameter, spherical, buoyant eggs, which after a brief planktonic phase of 5 to 10 h continue their development while adhering to a substrate. This short planktonic phase might explain why P. palpebratus only occurs around the Banda Islands, although similar habitats can be found elsewhere in the South Moluccan Sea. In addition to large ripe eggs, two size-classes of immature eggs were found in the ovaries. In the aquarium pairs of P. palpebratus tended to occupy a limited area but did not show aggressive territorial behaviour.     

Trophic and reproductive biochemistry of a deep-sea gelatinous octopus, Opisthoteuthis calypso

The present study provides a comprehensive analysis of the biochemical composition (proximate composition, amino acids and fatty acids in gonad, digestive gland and muscle) of the cirrate octopod, Opisthoteuthis calypso, collected off the coast of Portugal. Protein and energy contents of O. calypso were significantly lower (p < 0.05) than those observed for coastal octopods, suggesting that the gelatinous musculature of the cirrate octopus may not be a direct consequence of food limitation, but rather associated with reduced selective pressure for strong swimming ability in the deep sea. Moreover, principal component analyses of fatty acid (FA) data clearly separated coastal octopods from O. calypso. Saturated FA biomarkers (namely 16:0, 17:0, 18:0) and monounsaturated FA 22:1 suggest a higher phytodetrital influence on diet of O. calypso. A clear separation between groups was also observed with amino acid (AA) data. However, essential AA and non-essential AA designations (based on the flow of carbon through biochemical systems) do not necessarily provide an accurate picture of the origins of amino nitrogen in the deep-sea environment. Consequently, the interpretation of the present data is a challenging task but opens a new window of opportunity to unravel new trophic biomarkers in the deep sea.     

Observations on the embryonic development and energy source in the crab Xantho bidentatus

During development of Xantho bidentatus, ten stages of the eggs have been recognised. In this period the colour of the egg changes from black to pale white at the X stage of development. Total carbohydrate, glycogen, soluble protein, insoluble protein, total lipid, ash and water content have been determined in all ten stages of developing eggs. The utilisation of these compounds at various stages of development is discussed. The habitat of the crabs and its significance during development is also discussed. A constant decrease in the organic material as well as the dry weight of the egg was noticed with increased stage of development. Along with this a steady increase in the water and ash content was also observed. Glycogen was reduced during early days of development, whereas lipid was reduced during the latter stages of development. Both soluble and insoluble proteins exhibit conspicuous variations depending upon the stage of development.     

Synergistic effects of lead,salinity and temperature on embryonic development of the mussel Mytilus galloprovincialis

Combined effects of lead, salinity and temperature on the embryonic development of the mussel Mytilus galloprovincialis Lmk. were studied under laboratory conditions. The basic experimental design was a 4x6 factorial experiment using 4 lead concentrations (100, 250, 500 and 1000 ppb Pb²⁺) and 6 salinity levels (from 25 to 37.5 with 2.5 intervals). These factorial designs were carried out at three constant temperatures (150, 17.50 and 20°C). The statistical analysis indicated that salinity changes have more effect on the embryonic development than temperature. Optimal development was observed at 34.8 and 15.6°C, which is in accordance with observations in the field. The effect of lead was mininal in optimal salinity and temperature conditions. The deleterious effect of lead on the embryonic development was especially conspicuous at 20°C. Since in nature spawning occurs at temperatures inferior to 20°C, lead will probably not drastically decrease the potential recruitment of mussel spat in the littoral populations of the northern Adriatic Sea, where the salinity of the water is relatively stable. Under experimental conditions, lead caused a delay or inhibition of the embryonic development with the occurrence of a large number of abnormal larvae.     

Some observations on gametogenesis,larval development and substratum selection of the sea pen Ptilosarcus guerneyi

Ptilosarcus (Leioptilus) guerneyi (Gray) maintained in the laboratory, were observed to spawn in late March, 1972. Gametes, developed in the leaf proper, discharged through the mouths of feeding polyps and were fertilized externally in the sea water. The sea pen's eggs are 500 to 600 in diameter; a large female is capable of producing over 200,000 eggs in one season. A pear-shaped and free-swimming planula larva developed 4 days after fertilization, at a temperature of 12 °C. The larvae were ready to settle and metamorphose when 7 days old if favorable substratum was available, but would remain as planulae for at least 30 days if kept in glass dishes only. The 30-day-old larvae would metamorphose if a suitable substratum (coarse sand, for example) was presented. The larvae do not feed and, hence, development is lecithotrophic. Studies of histogenesis showed that metamorphosis greatly enhanced the rate of cellular differentiation. The high fecundity, lecithotrophic development, and the ability of substratum selection by the larvae explain the success of this species in maintaining a high-density population in many areas of sandy substratum in the shallow waters of Puget Sound (USA), despite the fact that it is preyed upon by 7 species of predators.     

Oxygen-dependent asynchrony of embryonic development in embryo masses of brachyuran crabs

Among brooding species, passive and active means to provide oxygen to embryos can be observed. Among passive oxygen providers, lower oxygen availability in the center than at the periphery of embryo masses seems to delay development of inner embryos. We investigated the differences in patterns of oxygen supply to the periphery and the center of embryo masses in two active oxygen providers, the brachyuran crabs Cancer setosus and Homalaspis plana, and evaluated the consequences on: (1) the proportion of time that early- and late-stage embryos were exposed to low or high oxygen partial pressure (PO₂), (2) oxygen consumption of the embryos from the center (inner) and the periphery (outer) of the embryo mass at those PO₂ levels that the embryos experience throughout development, and (3) development of inner and outer embryos. We found that oxygen availability in the embryo masses of brachyuran crabs exhibited dramatic contrasts between the periphery and the center during early development and that these differences decreased throughout embryonic development. These dissimilar patterns of oxygen availability produced differences in the proportion of the time that the embryos were exposed to high and low PO₂ levels throughout development. PO₂ affected oxygen consumption of the inner and outer embryos in the same fashion, but the oxygen demand of inner embryos was lower. Furthermore, development of inner embryos was delayed, in comparison to outer embryos of the same female. We suggest that the asynchrony in the development of inner embryos, in comparison to outer embryos, is due to oxygen limitation, since oxygen availability affects embryonic oxygen consumption. The differences between development of inner and outer embryos is relatively small, when compared to other marine invertebrates, probably because female crabs are able to adjust oxygen supply to the embryos according their needs, while passive oxygen providers are not. However, active oxygen provision may affect investment in reproduction. Our results could have important implications both on studies of larval development and survival and in understanding the life-history tradeoffs of aquatic invertebrates.     

Biannual spawning, rapid larval development and evidence of self-seeding for scleractinian corals at an isolated system of reefs

Some of the most important demographic parameters underlying the resilience of coral communities are determined by their patterns of reproduction. In this study, a variety of methods were used to investigate the patterns of spawning, larval development and dispersal for scleractinian corals at an isolated reef system off northwestern Australia. Two distinct periods of gamete maturation and multi-specific spawning occurred each year, during Spring and Autumn, in contrast to the single season of mass spawning described on most other reefs around Australia. The subsequent rates of embryogenesis and larval development were among the fastest described for corals, with pre-competency periods of approximately 3 days. Within 3 days of spawning, slicks of spawn and current drifters had dispersed up to 5 km, and up to 10 km after 6 days, while the times taken for drifters to travel between adjacent (>240 km) reef systems were similar to or greater than the upper competency periods of most coral larvae. Thus, under these conditions, the entire reef system, and to some extent the reefs within the system, may largely be self-seeded; rates of immigration from other systems are probably insufficient to rapidly increase the recovery of communities within years of a major disturbance. These results have implications for the management of isolated reef systems, highlighting the need to minimize local stressors and maximize community resilience to the increasing scale of disturbance and habitat fragmentation.     

In-situ observations of Baltic herring (Clupea harengus membras) spawning behaviour in the Askö-Landsort area,northern Baltic proper

In a Baltic herring (Clupea harengus membras L.) spawning ground study, carried out in summer 1982 in the northern Baltic proper, spawning herring were encountered on two line transects on June 14. The spawning behaviour of minor groups as well as of a large school was studied in situ by four divers for approximately three hours. No behavioural differences or systematic coordinated behaviour between the sexes could be detected. Most of the spawning took place in a limited depth interval, 0.5 to 4 m, and Ceramium tenuicorne was the substrate most used.     

Some observations of factors affecting embryonic and larval growth of Pollicipes polymerus (Cirripedia: Lepadomorpha) in vitro

Observations on the effects of several environmental conditions in embryonic and larval cultures are reported for the gooseneck barnacle Pollicipes polymerus (Sowerby, 1833). When the growth rates of control embryos kept in finger bowls at ambient-temperature sea water (ca. 12°–15°C) were compared to those of embryos grown under variously modified circumstances, it was found that growth rates were faster under conditions closely simulating the adult barnacle mantle-cavity (e.g. darkness and aeration). Addition of antimicrobial drugs and reduction of egg-mass size also promoted fast growth and development. Nauplii were fed several species of algae; only three species promoted growth to Stage V or older. Naupliar growth was fastest in larvae fed the combination of Prorocentrum micans/Platymonas sp. Larvae fed Prococentrum micans/pennate diatom grew slowest, and most larvae died at Naupliar Stage III. Interactions between algal species may have affected their nutritional value for Pollicipes polymerus nauplii. Less larval activity, slower growth rates, and higher mortality rates were observed in individuals given small amounts of food. The development of embryos and larvae in culture depends greatly on the culture conditions. These conditions should be described if comparison of timetables and envents are to be made between studies.     

Physiological and biochemical aspects of embryonic and larval development of the winter flounder Pseudopleuronectes americanus

Eggs and larvae of the winter flounder Pseudopleuronectes americanus Walbaum were hatched and raised in the laboratory under controlled conditions. Biochemical composition was measured during development and found to be similar to that of other species: 65 to 80 percent protein, 15 to 30 percent fat, and 0 to 5 percent carbohydrate. Ash content was 7 to 10 percent of dry weight. The chorion comprised more than half of the weight of an egg and the data suggested that it was possibly a source of nutrition to the developing embryo. The sequence of utilization appeared to be carbohydrate and then protein to hatching, lipid, mixed lipid and protein, the predominantly protein until feeding began. Carbohydrate was accumulated at first feeding and depleted when growth began. Protein and lipid were deposited in approximately constant proportions. Respiration rates of eggs were low, 0.002–0.015 l O₂ egg^-1 h^-1, but rose gradually from fertilization to hatching. Respiration rates of early larvae were from two to eight times that of eggs (0.033–0.131 l O₂ larva^-1 h^-1). Variation in larval respiration rates indicated a three-fold difference in rate according to level of activity. Eggs excreted ammonia at an increasing rate from fertilization to hatching. Larvae excreted ammonia, primary amines, and other unidentified organic nitrogenous substances. Rates of excretion and proportions of excretory products varied with stage of development. Primary amine excretion was variable and a major component in early stages. Ammonia-N excreted was two to 20 times primary amine N excreted. Unidentified substances were the predominant form of N excretion during early feeding. Ammonia accounted for most of the N excreted in older larvae. Early specific growth rates were 2.1 and 5.5%. Net caloric conversion and net and gross nitrogen efficiencies were low in first feeding larvae compared to adult fishes (32.2, 27.7, and 10.7% respectively).Contribution no. 5071 from the Woods Hole Oceanographic Institution     

Ovarian development and spawning in relation to the moult cycle in Northern krill, Meganyctiphanes norvegica (Crustacea: Euphausiacea), along a climatic gradient

Adaptive processes linked to reproduction were studied comparatively for three populations of Northern krill, Meganyctiphanes norvegica (M. Sars, 1857), sampled during winter and summer cruises in the Clyde Sea (W Scotland), the Kattegat (E Denmark), and the Ligurian Sea (NW Mediterranean). The aim was to investigate the functional relationship between egg production and moulting under contrasted climatic and environmental conditions. A staging system for female sexual development established for live krill was complemented by a histological study of the ovary at various developmental steps. During the reproductive season, all adult female krill were engaged in cyclical egg production. During experiments, female krill released one batch of mature oocytes in one or two spawning events. The ovary of postspawn female krill still contained developing oocytes for another egg batch. In the non-reproductive period, all female krill had a resting ovary. Ovarian structure and pattern of egg production were identical in the three populations, but seasonal timing of egg production was different. The model proposed for the Ligurian population of the annual cycle of ovarian development can be extended to the other two populations, taking into account the seasonal characteristics of each site. Random field samples were staged simultaneously for moult cycle and for sexual development. Moult stages and the seasonal variation of the intermoult period were studied for the Kattegat population using multi-year data and compared to data obtained during summer/winter cruises in the Clyde and the Ligurian Sea. At the three sites, intermoult period was shorter and temperature-dependent during the reproductive period, concurrent with the season of greatest food availability. During most of the year and the period of sexual rest, moulting activity was reduced. The relationship between spawning and the moult cycle was studied comparatively for the three populations. Eggs were released during the premoult phase of a “spawning moult cycle”, in one or two spawnings associated with apolysis and Moult Stage D1, respectively. Yolk accumulation for the next egg batch was completed during an alternating “vitellogenic moult cycle”. A model for the timing of cyclical egg production in relation to moulting, as proposed for the Kattegat, can be extended to the other populations, taking into account intermoult period variation with temperature. Temperature appeared to be the principal environmental factor controlling growth (through moulting) and egg production during the reproductive season, in connection with favourable trophic conditions. Received: 22 December 1997 / Accepted: 29 August 1998     

Laboratory studies of the marine copepod Centropages typicus: egg production and development rates

Egg production and development rates of Centropages typicus (Krøyer) were studied in the laboratory under carying food and temperature conditions. Egg production rates in the laboratory ranged from 0 to 124 eggs female^-1 d^-1 and increased with food concentration up to a critical food concentration (P_c) above which egg production was constant. Egg production rates were influenced by temperature, with more eggs being produced at 15°C than at 10°C. Thalassiosira weisflogii and Prorocentrum micans were determined to be equally capable of supporting egg production at concentrations above P_c at 15°C. Rate of egg production was independent of adult female size when food and temperature were constant. Egg production rates of freshly captured females ranged from 0 to 188 eggs female^-1 d^-1 and were higher in April and May than in June or July. Hatching rates of eggs increased with increased temperature; 95% of the eggs at 15°C hatched within 48 h, while only 8% of the eggs at 10°C hatched within 48 h. Development rates, determined at 10°C in excess concentrations of T. weisflogii, were 23.0 d from egg release to copepodid state I, 27.0 d to stage II, 29.5 d to stage III, 32.2 d to stage IV, 38.5 d to stage V and 49 d to adulthood based on the average time required for 50% of the organisms in an experiment to attain a given stage. Adult males were usually observed 2 to 4 d before adult females, and therefore have a slightly faster rate of development. The effects of temperature, food type and food concentration on egg production and the seasonal appearances of diatoms in the New York Bight may account for the observed seasonal cycles in abundance of C. typicus in these coastal waters.     

Seasonal observations on the biometry and development in Copepodite Stage V of Calanus finmarchicus from the Firth of Clyde

Relationships between linear dimensions of casts produced at the final moult in the laboratory and definitive sex confirm that Calanus finmarchicus (Gunn.) is sexually dimorphic in Copepodite Stage V. Two forms of Stage V copepodite were present at all times of the year in the Firth of Clyde, Scotland. Except in early spring, when size was similar, these were consistently distinguished by the prosome length. The small form moulted to adult females and the large form to adults of either sex. Relative to the metasome, the length of the cephalosome was greater in the large than small form of Stage V copepodite. This, to an increased extent, distinguished males from all females in the adult. Differences between females of the large and those of the small form were similar in the Stage V copepodite and adult. Feminization in the large form entailed an arrest in the differentiation of secondary male sex-characters. Most morphological females moulting from both forms of Stage V copepodite possessed an apparently normal ovary, with eggs extending into oviducal diverticula. In the case of the large form, however, eggs were occasionally absent from diverticula. The gonad in two specimens resembled an immature testis. In males, a normal testis was always present. Differentiation into the adult male entailed the internal suppression of feminizing factors. On average, within collections, females of the large form moulted later than the small form and definitive males. A similarly consistent difference between females of the small form and males was absent. Moulting times in the large form of Stage V copepodite depended on the presence or not of feminization. Differentiation into the adult male required the environmental stimulation of development. Comparisons with moulting patterns in the sea suggest a mechanism that regulates the recruitment of males to the adult.