The gametogenic cycle of<Emphasis Type="Italic"> Octopus vulgaris</Emphasis> (Mollusca: Cephalopoda) as observed on the Atlantic coast of Andalusia (south of Spain)

The present study reports the gametogenic cycle of the common octopus, Octopus vulgaris Cuvier, 1797 in the Atlantic coast of Andalusia between May 2001 and May 2002. A total of 673 specimens of total weight ranging from 30 to 3,732 g were analysed. The gonad development was defined by means of histological methods and gonadosomatic indexes. Four gametogenic stages were identified in males (I immature, II maturing, III mature, IV spawning) and in females (I immature, II maturing, III pre-spawning, IV spawning). The sex ratio obtained was not significantly different from 1:1 (P>0.01). Mature males were found during the whole year, September being the month when the highest rate of sexually active individuals occurred (48%). In females, the gametogenic cycle commenced in December and ended in November, and spawning occurred between April and October, with two maximum peaks in August and September (48% in each). Size at sexual maturity was 850 g of total body weight for males and 1,250 g of total body weight for females. Males mature earlier than females.Communicated by S.A. Poulet, Roscoff     

The effects of symbiotic state on heterotrophic feeding in the temperate sea anemone <Emphasis Type="Italic">Anthopleura elegantissima</Emphasis>

The temperate sea anemone Anthopleura elegantissima is facultatively symbiotic with unicellular algae. Symbiotic A. elegantissima can supplement heterotrophic feeding with excess photosynthate from their algal partners, while asymbiotic individuals must rely solely on heterotrophy. A. elegantissima individuals were collected from Swirl Rocks, Washington (48°25′6″ N, 122°50′58″ W) in July 2010, and prey capture and feeding characteristics were measured to determine whether asymbiotic individuals are more efficient predators. Feeding abilities were then measured again after a 3-week exposure to full sunlight or shaded conditions. Freshly collected asymbiotic anemones had larger nematocysts, but symbiotic individuals showed greater nematocyte sensitivity. Sunlight enhanced digestion and reduced cnida density in all anemones regardless of symbiotic state. Results suggest that the phototropic potential of A. elegantissima, as influenced by symbiotic condition, has little effect on heterotrophic capacity. The anemones appear to maximize heterotrophic energy input independent of the presence or identity of their algal symbionts.     

Annual reproductive cycle of the green sea urchin, Strongylocentrotus droebachiensis, in differing habitats in Nova Scotia, Canada

We monitored the reproductive cycle of Strongylocentrotus droebachiensis (OF Müller) between April 1993 and August 1995 in kelp beds, barren grounds and grazing fronts at both a wave-exposed and a sheltered site along the Atlantic coast of Nova Scotia. Gonad index and histological analyses showed that S. droebachiensis has an annual reproductive cycle that is synchronous across sites and habitats, and between females and males. Spawning occurs in March/April of each year but a small proportion of sea urchins in the study populations also spawned in fall 1995. During most of the year, sea urchins in kelp beds and grazing fronts have a higher gonad index than those in barren grounds. Gonad indices also tended to be higher at the wave-exposed than the sheltered site. Interannual variability in peak gonad index was significant in the barren grounds at the wave-exposed site and in the grazing front at the sheltered site. The gametogenic cycle is characterized by six stages based on the abundance of nutritive and germinal/gametic cells. Nutritive phagocytes are abundant after spawning and replaced by increasing numbers of germinal and gametic cells as the gametogenic cycle progresses. The temporal patterns of abundance of each cell type were similar among habitats indicating that the gonads were qualitatively similar despite large differences in gonadal mass. The quantity of gut contents (ratio of food volume to body volume) was similar among habitats, but the quality (percentage of organic material) tended to be higher in kelp beds and grazing fronts than in barren grounds suggesting that differences in gonad index of S. droebachiensis in different habitats are related to differences in diet. The high density of sea urchins in grazing fronts combined with their high fecundity suggests that they make the greatest contribution, per unit area, to the overall larval pool. Received: 20 May 1997 / Accepted: 21 January 1998     

The reproductive cycle of <Emphasis Type="Italic">Scolelepis goodbodyi</Emphasis> (Polychaeta,Spionidae)

The aim of this study was to examine the reproductive and gametogenic cycle of the spionid polychaete Scolelepis goodbodyi (Jones). Every 15 days, in the upper level of the intertidal zone on Barequeçaba Beach in southeastern Brazil, a 100 m² area was delimited and within it three points were selected and sampled using a core sampler 0.01 m² in area × 0.2 m long. Each sample was divided into three sub-samples: surface (2 cm) and middle (8 cm), which were sieved through 0.5, 0.25 and 0.125 mm mesh sieves; and lower (10 cm), sieved through 1.0 and 0.5 mm mesh sieves. Males, females and undetermined individuals were separated, and the width of the third setiger was measured. The gonadal development of each individual was classified based on external features of the gametogenic setigers. Ten to 20 adults from each sampling period were classified according to their gonadal development stage and then analyzed histologically for gametogenic studies, for comparison with the former classification. Some of the ripe females were used for fecundity evaluation. The diameters of the oocytes were measured prior to spawning. Eight hundred and twenty-eight females (29.70%), 848 males (30.42%) and 1,112 undetermined individuals (39.89%) were analyzed. The sex ratio did not differ significantly from unity (χ² = 0.24; df = 1; P < 0.05). Males and females were more abundant in May 2002 (ca. 80%) and between November 2002 and May 2003 (between 77 and 90%); the proportion of undetermined individuals did not represent recruitment alone, but also a high incidence of adults with recovering gonads. Fecundity varied from 134 oocytes in a 0.54 mm W3 individual to 289 oocytes in a 0.4 mm W3 individual, with a mean of 220 (SD = 57) oocytes per female and 19 (SD = 5) oocytes per gametogenic setiger. Oocytes were ellipsoid, and ranged in size from 150 × 80 (to 220 × 120 μm (mean ± SD = 173 × 113 ± 15 × 11 μm). Neither a well-defined gametogenic cycle nor synchrony in the reproductive period was observed, because many gonadal development stages occurred in each month. Nevertheless, peaks of maturity were observed between October–December 2002 and April–May 2003.     

Reproduction and annual gametogenic cycle in Nephtys hombergii and N. caeca (Polychaeta: Nephtyidae)

The gametogenic cycles of Nephtys hombergii Audouin et M. Edw. and N. caeca Fabricius are described. Both species are polytelic (=discrete, iteroparous) and breed in their second and subsequent years. They spawn in May in the Tyne estuary. After spawning, N. hombergii retains a large number of gametocytes, which are ultimately resorbed. A new generation of gametocytes is proliferated by September and mature gametes accumulate during the winter months. Gametic development is not well synchronised.     

Reproductive periodicity of the scleractinian coral Lophelia pertusa from the Trondheim Fjord, Norway

Between 2002 and 2008, samples of the cold-water scleractinian coral Lophelia pertusa were collected from the Trondheim Fjord in Norway to examine reproductive periodicity. Collections were made from three locations: Tautra, (63°35.36′N, 10°31.23′E at 40–70 m), Stokkbergneset (63°28.18′N, 09°54.73′E at 110–500 m), and Røberg (63°28.88′N, 09°59.50′E at 250 m). Populations of L. pertusa from the Trondheim Fjord initiated oogenesis in January and spawning occurred from late January to early March the following year. Gametogenic cycles of the female L. pertusa samples overlapped by approximately 2 months, with oogonia visible in January, but this was not evident in the males. This paper provides the most complete gametogenic cycle to date and spawning observations for this important structure-forming species. The results from fjord populations are compared with published and preliminary data from other regions and are discussed in the context of regional differences in physical and biological variables, particularly food supply. Differences in gametogenic cycles within a single species provide a rare opportunity (especially in deep-sea species) to examine potential drivers of reproduction.     

Reproduction in the diadematoid sea urchin Centrostephanus rodgersii in contrasting habitats along the coast of New South Wales, Australia

Reproduction in the sea urchin Centrostephanus rodgersii was examined in two types of habitats (“barrens”, i.e. habitats characterised by the high crustose coralline algal cover typical of urchin-barren grounds, and by the absence of macroalgae; and “fringe”, i.e. habitats characterised by a high macroalgal biomass and few C. rodgersii) at four locations in New South Wales. The four locations: the Solitary Islands, Sydney, Ulladulla and Eden, span the distribution of C.␣rodgersii from the subtropics at its northern limit to temperate waters near its southern limit. Histology and estimates of gonad retrieval rate (GRR) from January 1994 to October 1995 indicated that reproduction was synchronous at all locations. An increase in the tempo of gametogenesis in May and onset of spawning in June at all locations is consistent with entrainment in response to exogenous factors. Over the range studied, C. rodgersii experienced relatively similar daylength cycles and contrasting sea-temperature cycles. Short days and lunar conditions coinciding with the solstice appear likely proximate cues for the onset of spawning. The major difference in reproduction among locations was in the duration of spawning. In the southern parts of its range breeding occurred over a 5 to 6 mo period, whereas at the Solitary Islands it lasted ≃1 mo. At most locations the GRRs were significantly higher in the fringe habitat than in the barrens habitat. The lower reproductive output of urchins in the barrens habitat was attributed to the food-poor conditions typical of this habitat. The developing fishery for C. rodgersii is likely to be most effective from March to early May. Urchins from barrens areas may not provide sufficient yield to warrant harvesting. Received: 29 October 1997 / Accepted: 18 May 1998     

Sea anemones and brittle stars: unexpected predatory interactions during induced in situ oxygen crises

During oxygen crises, benthic faunas exhibit a series of behavioural patterns that reflect the duration and severity of the event. During artificially induced oxygen deficiencies at 24 m depth in the Northern Adriatic Sea, we photographically documented predation by the sea anemones Cereus pedunculatus (Pennant, 1777) and Calliactis parasitica (Couch, 1842) on the brittle star Ophiothrix quinquemaculata (DelleChiaje, 1828). Five predatory events were recorded with four anemones during nine deployments totalling 817 h of observation. Under near-anoxic conditions, individuals of both actinians made contact with, pulled in and consumed the brittle stars. The duration of each predatory event was 1.5–7.5 h. In three of the five events, brittle star remains were regurgitated after an additional 2.0–12.5 h of digestion by the anemones. Our time-lapse sequences demonstrate that oxygen deficiency, beyond eliciting a series of specific behaviours in members of each species, also promotes previously unobserved interspecific interactions. Our results show that sea anemones are not only highly resistant to anoxia, but may also benefit by taking advantage of prey that are more vulnerable to anoxic conditions.     

Effect of feeding regime and irradiance on the photophysiology of the symbiotic sea anemone Aiptasia pulchella

To determine how the animal and algal components of the symbiotic sea anemone Aiptasia pulchella respond to changes in food availability and culture irradiance, sea anemones from a single clone were maintained at four irradiance levels (320, 185, 115, and 45 E m^-2 s^-1) and either starved or fed for 5 wk. Changes in protein biomass of sea anemones maintained under these conditions were not related to the productivity of zooxanthellae, since the protein biomass of fed A. pulchella decreased with increase in irradiance and there was no difference in protein biomass among starved sea anemones at the four irradiance levels. Except for the starved high-light sea anemones, the density of symbiotic zooxanthellae was independent of culture irradiance within both starved and fed. A. pulchella. Starved sea anemones contained over twice the density of zooxanthellae as fed sea anemones. Within both starved and fed individuals, chlorophyll per zooxanthella increased with decreasing culture irradiance while algal size remained constant (in fed sea anemones) at about 8.80 m diameter. Chlorophyll a: c ₂ ratios of zooxanthellae increased with decreasing culture irradiance in zooxanthellae from starved sea anemones but remained constant in zooxanthellae from fed sea anemones. As estimated from mitotic index data, the in situ growth rates of zooxanthellae averaged 0.007 d^-1 and did not vary with irradiance or feeding regime. Photosynthesis-irradiance (P-I) responses of fed A. pulchella indicated an increase in photosynthetic efficiency with decreasing culture irradiance. But there was no consistent pattern in photosynthetic capacity with culture irradiance. Respiration rates of fed sea anemones also did not vary in relation to culture irradiance. The parameter I _k , defined as the irradiance at which light-saturated rates of photosynthesis are first attained, was the only parameter from the P-I curves which increased linearly with increasing culture irradiance. The daily ratio of net photosynthesis to respiration for A. pulchella ranged from 1.6 to 2.8 for sea anemones maintained at the three higher irradiances, but was negative for those maintained at 45 E m^-2 s^-1. Since the final protein biomass was greatest for sea anemones maintained at the lowest irradiance, these results indicate that sea anemone growth cannot be directly related to productivity of zooxanthellae in this symbiotic association.     

Physiological energetics of the intertidal sea anemone Anthopleura elegantissima

High-intertidal (H) individuals of the sea anemone Anthopleura elegantissima (Brandt) are exposed aerially up to 18 h each day, unlike low-intertidal (L) individuals which may be continuously immersed over many days. Thus, H anemones experience shorter feeding periods compared to L anemones. From 1980 to 1982, H and L anemones were observed and collected at the mouth of Bodega Harbor in North Central California (USA) to determine whether any physiological adaptations mitigate the energetic effects of reduced feeding time in H anemones. Weight of prey in coelenterons of H anemones was three times more than that of L anemones following a single immersion period. This difference is not due to slower digestion rates in H anemones. Prey residence time in coelenterons (4h) was equivalent in both groups. Different prey weights imply that ingestion rates were greater in H individuals. However, all anemones had similar weight-specific feeding-surface areas. Different prey-capture rates result from increased receptivity to prey in H anemones, rather than from increases in feeding surface. Absorption efficiency was inversely related to ration size in anemones from both shore positions. H individuals absorbed food more efficiently than L individuals fed equivalent rations. Ration, not exposure conditions, affected absorption efficiency. Daily growth rates were 1.5 to 1.8% and 1.2 to 1.4% of dry body weight in H and L anemones fed large rations (4.0 to 5.6% of dry body weight), respectively. H anemones fed smaller daily rations, approximating amounts of zooplankton captured naturally (1% of anemone dry weight), had higher growth rates and growth efficiencies than L anemones, which lost mass. Higher growth rates in H anemones, which are supported by higher prey-capture rates, result in attainment of minimum body size for reproduction in a relatively short period of time despite reduction in time available for feeding, thus improving relative fitness of these anemones in the upper intertidal zone.     

Synchronous aggregated pseudo-copulation of the sea star <Emphasis Type="Italic">Archaster angulatus</Emphasis> Müller &; Troschel, 1842 (Echinodermata: Asteroidea) and its reproductive cycle in south-western Australia

Mass individual pairing or pseudo-copulation of the sea star Archaster angulatus was observed in November and December 2009, suggesting a late-spring/summer spawning period for this species on the west coast of Australia. Detailed measurements were made on the second of these occasions. Density of sea stars was 1.11 per m² and 68.5% were in mating pairs. Copulating pairs were mostly male on female, occasionally male on male. There was no difference in size between males and females in mating pairs. No evidence was found to indicate mating and spawning is coincident with lunar or tidal cycles. Females outnumbered males by more than 20%, but the difference in sex ratio was not statistically significant. Analysis of the reproductive cycle revealed that gonad indices reached their peak in October and declined from then until January. Histological sections of gonads confirmed that sea stars are in peak reproductive condition in October and November and are fully spent by January. Males have a much lower (ca. 1/3rd) gonad index than females when each are in peak reproductive condition, the second lowest recorded for any sea star. Pyloric caecae indices showed little annual variation and monthly averages of just 3–4% are among the lowest ever recorded for an asteroid. It is suggested that these characteristics are associated with the copulatory behaviour of the deposit feeding A. angulatus, enabling the species to maintain a high level of fertilisation success while also minimising the allocation of energy to gonad development in habitats with low or variable food availability. However, it is still difficult to explain why a species which ensures a high level of fertilisation by pseudo-copulation also does this en masse and synchronously. One hypothesis is that competition for males and the benefits of having eggs fertilised by multiple males favours both synchrony and aggregation.     

Synchronous spawnings of 105 scleractinian coral species on the Great Barrier Reef

Following observations of mass spawning of hermatypic corals on the Great Barrier Reef in 1981 and 1982, spawning dates were successfully predicted and documented at five reefs on the Central and Northern Great Barrier Reef in 1983. During the predicted times, 105 species from 36 genera and 11 families were observed to spawn. Of these, 15 species were shown to have an annual gametogenic cycle. All but two of the species observed during mass spawnings shed gametes which underwent external fertilization and development. Synchronous spawning was observed both within and between the five reefs studied, which were separated by as much as 5° of latitude (500 km) or almost a quarter of the length of the Great Barrier Reef. The mass spawning of corals took place on only a few nights of the year, between the full and lastquarter moon in late spring. Maturation of gametes coincided with rapidly rising spring sea temperatures. Lunar and diel cycles may provide cues for the synchronization of gamete release in these species. The hour and night on which the greatest number of species and individuals spawned coincided with low-amplitude tides. Multispecific synchronous spawning, or mass spawning, of scleractinian and some alcyonacean corals represents a phenomenon which is, so far, unique in both marine and terrestrial communities.     

Contrasts between spawning times of <Emphasis Type="Italic">Anguilla</Emphasis> species estimated from larval sampling at sea and from otolith analysis of recruiting glass eels

This study reviewed literature on spawning times for three north temperate species of anguillid eels estimated by sampling for small leptocephali (larvae) at sea and for several temperate and tropical species by back-calculating from putative daily ages derived from otolith increment analysis of glass eels that recruited to coastal waters. Estimates from otoliths of European eels, Anguilla anguilla, American eels, Anguilla rostrata, and Japanese eels, Anguilla japonica, imply much more protracted spawning seasons than are indicated by sampling at sea during various times of year. European eels are inferred to spawn year-round from otolith analysis, but the smallest, recently hatched leptocephali are found only in late winter and spring. From otoliths, the spawning times of these three species are all estimated to occur much later in the year than when small leptocephali are found at sea, indicating that ages appear to be underestimated. For these and other temperate and tropical eels, there are inconsistencies in assigned ages among various studies, which are most extreme for the European eel. This species has the longest larval migration and often has an opaque zone in the glass eels’ otoliths where it is difficult to discern growth increments. These inconsistencies suggest that interpretation of otolith growth increments is incorrect at least in some studies, and the apparently consistent mismatch between otolith and sea-sampling studies suggests that increments may not always be formed at some period during the unusual early life history of anguillids. Because daily increments may be formed in eels during most of their early life history, future research is needed to determine the cause of the mismatch of glass eel aging studies and the apparent spawning times of eels offshore. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.

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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.     

Reproductive energetics,endocrinology and population dynamics of Nephtys caeca and N. hombergi

An investigation of gametogenetic cycles and spawning success in populations of Nephtys caeca and N. hombergi from the River Tyne estuary, North Shields, England, in relation to production of gonadotrophic (GH) and spawning hormones (SH) has been made, based on data collected from 1978–1984. The data show that a consistent pattern of gametogenesis and efficient spawning occurs in populations of N. caeca in north-east England, but that the reproductive cycle of N. hombergi is erratic. Premature oosorption and gametogenic failure in this latter species is associated with a low level of GH production. Following high levels of GH production, gravid females sometimes fail to spawn owing to the non-release of SH. The reproductive energetics and reproductive success of N. caeca and N. hombergi are described in relation to the observed patterns of endocrine activity and the consequent changes in population structure.     

The role of fecundity regulation and abortive maturation in the reproductive strategy of Norwegian spring-spawning herring (<Emphasis Type="Italic">Clupea harengus</Emphasis>)

This study investigates the reproductive strategy, an important component in the estimation of stock reproductive potential, in Norwegian spring-spawning (NSS) herring (Clupea harengus), an iteroparous, extreme capital spawner, through the estimation of fecundity over a period of 3 years including two complete maturation cycles and three spawning seasons. NSS herring have an ‘optimistic’ strategy, with almost all adult herring caught in August being in the vitellogenic stage of ovary development, despite overwintering energy levels not being determined at this time. Fecundity in the summer, i.e., more than half a year before spawning in spring (February–April), was also much higher than could be supported by an individual’s concurrent energy levels. Consequently, fecundity was later reduced through atresia with the majority of this occurring before overwintering. The total reduction and the length of the time period in which the reduction took place appeared to vary between years. During the spawning season, atresia was mostly prevalent in small first-time spawners <180 g and several individuals aborted ovary development at this time. Final fecundity varied between years with a difference of up to 18% and was linked to annual variations in condition. In conclusion, this extensive field study has demonstrated that each individual herring can display a suite of size-specific reproductive tactics to fine-tune oocyte production in response to fluctuating levels of planktonic prey.     

Growth and survivorship in a tidal-flat population of the bivalve Mercenaria mercenaria from Southampton water

A monthly sampling programme was conducted to investigate aspects of the biology of a population of Mercenaria mercenaria (L.) Smooth growth curves were constructed for each year class, and the factors affecting growth rate were considered. Seasonal cycles in flesh weight and calorific content appear to be related to gonal proliferation and subsequent spawning. Recruitment is sporadic and probably depends on the spawning of upstream populations, where conditions are more favourable for larval development. Survivorship curves were constructed for each year class; predation by crabs and gulls appears to account for a large proportion of the observed mortality.     

Effects of starvation,and light and dark on the energy metabolism of symbiotic and aposymbiotic sea anemones,Anthopleura elegantissima

Rates of oxygen and carbon-dioxide exhange were measured in symbiotic and aposymbiotic specimens of the sea anemone Anthopleura elegantissima while fed and starved under light or dark conditions. Respiratory quotients indicated that fed anemones switched from a carbohydrate to a fat catabolism when starved, with the exception that symbiotic individuals starved in the light showed a pronounced carbohydrate catabolism for over 1 month. The source of the carbohydrate was probably photosynthate translocated by the dinoflagellate Symbiodinium (=Gymnodinium) microadriaticum (Freudenthal) living in the anemones' tissues. The starved symbiotic anemones maintained in the light had lipid levels not significantly different from fed controls and 44 to 61% higher than starved aposymbiotic anemones after 1 month. Thus, the quality and quantity of the metabolic flux from the symbionts to the sea anemone were sufficient to conserve the host's lipid reserves.