Maturation,epitoky and regeneration in the polychaete Eunice siciliensis under field and laboratory conditions

In the gonochoric polychaetous annelid Eunice siciliensis Grube, germ cells only develop in the posterior segments of the worm; these segments subsequently differentiate into an epitokous portion separating at maturity from the infertile body part. As in the Pacific and Atlantic palolo worms, the epitokous portions die some hours after spawning. Laboratory investigations revealed that the anterior parts of spawned worms survive and regenerate genital segments. After a period of at least 8 months, paired gonads are developed in these segments. Gonia proliferating later from the germinal tissue are released from gonadal sacs into the coelomal cavities, where gamete development occurs. A reproductive epitokous portion does not mature and separate until the second year of cultivation; it is followed by caudal regeneration. These facts and the observation that only about 50% of the individuals obtained from samples of coralligène were sexually differentiated, led to the conclusion that E. siciliensis probably has a bi-annual reproductive cycle. Caudal regeneration does not depend on the presence of a prostomia hormone. The ability to form posterior regenerates is inherent to the parapodial segments of all body regions, except for mature genital segments. This is also valid for the formation of morphologically different secondary prostomia. However, for initiation and further promotion of gametogenesis and epitokous development, at least one gonadotropic factor originating from the prostomium is required. Neurosecretory cells within the brain suggest this factor(s) to be of neuro-endocrine nature. Specimens developing secondary prostomia were found both under field and laboratory conditions. In several cases, sexual development took place, and even epitokous portions were separated from such individuals. From this it appears that the gonadotropic functions are re-established by the regenerated prostomia in order to ensure reproduction in individuals accidentally deprived of the anterior end. The possible rôle of temperature and light as environmental factors involved in the control of normal and regenerative growth, of sexual development and reproduction is discussed.     

Nachweis der epitokie als fortpflanzungsmodus des polychaeten Eunice siciliensis (Polychaeta: Annelida)

Eunice siciliensis (Grube)¹ is a sedentary polychaete with separate sexes; its germ cells develop only in the posterior part of the body. In March and April, females with whitish or even dark-green oocytes, males with only few spermatogonia, and individuals with no germ cells at all were discovered in the “coralligène” of the Banyuls region (European Mediterranean Sea). In august, besides immature and sexually non-differentiated worms, mature individuals of both sexes were captured: females with dark bluish-green oocytes of diameters up to 250μm, and males with milky genital segments containing spermatozoa. Soon after isolation from the substrate, these individuals performed characteristic movements with their posterior body parts, which then autotomized, and released ova and spermatozoa. Evidence is given that the anterior, atokous body parts survive in their tubes in the “coralligène” after separation from the epitokous portions, and regenerate new genetal segments. The development of gametes observed in these regenerated segments indicates the ability of E. siciliensis to reproduce more than once during its lifetime. Not only caudal, but also prostomial regeneration was observed; the formation of a (morphologically different) secondary prostomium appears to be essential for any development of germ cells in worm fragments. E. siciliensis exhibits surprising similarities in habits and in formation of an epitokous form with Eunice viridis (Gray), the “palolo” worm of the South Pacific Ocean, but nevertheless, lacks the prominent criteria of this famous species: the ventral eye spots on each genital segment in males and females, and the paired areas with brown pigmentation on the ventral side of each epitokous segment in the males.     

Reproduction, larval development and functional relationships of the burrowing, spionid polychaete Dipolydora armata with the calcareous hydrozoan Millepora complanata

Dipolydora armata (Langerhans, 1880) is a small (4 to 5 mm) spionid polychaete found burrowing in the calcareous hydrozoan Millepora complanata Lamarck, 1816, on coral reefs at Barbados, West Indies. It excavates complex networks of interconnecting burrows and forms aggregations of worms in cavities within branches of the coral. Adult worms have a mixed feeding mode (suspension feeding and deposit feeding). Size–frequency distributions of worms in branch samples suggest that they mature in a single year and that reproduction occurs throughout the year. Burrow openings on the surface of the coral develop distinctive, erect spines caused by combined growth of worm tubes and host tissue. Millepore zooids were absent in the vicinity of tube openings and on spines, and thus the potential feeding surface of the coral will be reduced in heavily colonized branches. Burrows and openings were densest at the bases of millepore branches where weakening of the skeleton would be expected to occur. The absence of openings near the branch tips suggests difficulty in larval settlement there, amongst stinging zooids. Reproduction␣and larval development of the worms were examined, and a sequence of larval stages from one to 20 segments and a juvenile stage of 22 segments are described. Eggs are deposited in brood sacs attached to the burrow wall, and the larvae feed upon nurse eggs (adelphophagy). The presence of larvae and juveniles occurring free in the burrows suggests that larval development may be completed within the host coral as an alternative or in addition to a planktonic larval phase. Lack of provisional larval setae, early development of adult capillary setae, production of special spermatophores and a protracted breeding cycle in D. armata are all traits which would favour complete development within the host skeleton. Received: 6 March 1997 / Accepted: 25 October 1997     

Moon orientation in sandhoppers: effects of lighting treatments on the persistence of orientation ability

The moon orientation rhythm persists in sandhoppers removed from environmental entrainments relative to moon periodicity. In order to investigate the underlying mechanism, the crustacean amphipods Talorchestia capensis and Talitrus saltator were collected at new moon and exposed in the laboratory to an artificial light during the day, but to a variety of lighting treatments during the night: (1) continuous dark, (2) dim light in phase with the natural moon, (3) continuous dim light. The animals were tested under the moon 9–21 days later by using a new type of arena. The ability to orientate in a direction perpendicular to the shore persisted in sandhoppers under treatments 1 and 2, but not under treatment 3. A disturbance due to a phototactic tendency in Talorchestia was also observed in animals captured on the day of the test when exposed to unnatural lighting, but not when they were kept under natural light. The present findings show that the timing mechanism allowing compensation for changes in the moon’s position also persists in animals that have been long removed from entraining factors. The dependence of this orientation ability on nocturnal lighting and the disturbing effect of sudden changes in light intensity support the idea that the phasal lighting of the moon, and perhaps sunrise and sunset act as resetting factors for the moon’s orientation rhythm.     

Seasonality,lunar periodicity of settlement and microhabitat association of juvenile humpback red snapper <Emphasis Type="Italic">Lutjanus gibbus</Emphasis> (Lutjanidae) in an Okinawan coral reef

Snappers (Lutjanidae) are one of the important fisheries resources in tropical and subtropical waters. However, there have been few studies clarifying the ecology of newly settled juvenile snappers. The aims of the present study were to clarify the seasonality and lunar periodicity of the larval settlement, and the microhabitat association of newly settled juveniles for Lutjanus gibbus in an Okinawan coral reef. Fifteen coral patches were chosen and underwater visual surveys were conducted during a 2-year period. The larval settlement of the species mainly occurred between May and October during the 2-year survey period. In terms of lunar periodicity of the settlement, the newly settled juveniles were mainly found between the last quarter moon and new moon during the main settlement season. Back calculations of settlement dates based on otolith microstructure showed that most juveniles settled during the last quarter moon and new moon. Stepwise multiple regression analyses revealed that the newly settled juveniles were mainly found on coral patches with a large number of holes on the top of the coral patch and this tendency was mainly consistent during the study period. The volume of holes at the base of coral patches had also a positive effect on the abundance of newly settled juveniles in some cases. The results of the present study suggest that larval settlement has a strong seasonal and lunar periodicity, and newly settled juveniles select coral patches with available refuge spaces.     

Diel and lunar patterns of reproduction in the Caribbean and Pacific sergeant major damselfishes Abudefduf saxatilis and A. troschelii

Diel and lunar reproductive patterns of the Caribbean and Pacific sergeant major damselfishes Abudefduf saxatilis and A. troschelii respectively, were studied i the San Blas and Northern Perls Islands of Panama in 1981 and 1982. These are sister-species that have diverged since the uplift of the isthmus of Panama. Caribbean A. saxatilis females typically complete spawning in the nests of males earlier in the morning than do Pacific A. troschelii females. Spawning is rarely observed during the afternoon. Hatching occurs four or five days after spawning in the Caribbean species, and after four days in the Pacific species. In both, fry hatch during the hour following sunset. There also are differences in lunar patterns of spawning between the two species. The Caribbean A. saxatilis spawns throughout the month, exhibiting no lunar periodicity. In the Pacific A. troschelii spawns within nine days on either side of new moon. During the wet season, reproductive activity at a site spans three or four days, but is extended to eight or more days during dry season upwelling in the Bay of Panama. Thus, reproductive synchrony is much greater within than between colonies of the Pacific sergeant major, particularly in the wet season. Reproductive synchrony in the Pacific sergeant major might be beneficial because fry hatching at dusk encounter favorable tides for transport away from reef-based predators around new moon, or because reproductive synchrony reduces the risk of predation on unhatched embryos in nests guarded by males. Tidal amplitude and the rate of predation on embryos in nests are greater in the Pacific.     

Occurrence and biological role of light-induced chromatophore displacements in seaweeds

The occurrence of light-induced chromatophore displacements and concomitant transmittance changes in marine algae was investigated by microscope and photometrically with an automated recording microphotometer system; 16 brown, 6 green and 20 red algae were studied. In most of the brown algae, both phaeoplast displacements and transmittance changes were found. In some red algae which are frequently exposed to direct sun light during emergence at low tide, light-induced transmittance changes were measured, but they could not unequivocally be correlated with changes in the position of rhodoplasts. Among green algae, only Ulva lactuca shows chloroplast displacements which, however, follow circadian rhythms and are consequently not light-induced in the strict sense. The dose-response curves of light-induced chromatophore displacements were measured in the following Fucus and Laminaria species: F. spiralis, F. vesiculosus, F. serratus, L. digitata, L. saccharina and L. hyperborea. While in Fucus species correlations between light-induced transmittance changes and zonation of the intertidal area seem to exist, no significant differences have been found in the Laminaria species. The physiological role and ecological importance of light-induced chromatophore displacements for seaweeds living in the intertidal belt are discussed.     

Cyclical mucous sheet formation on poritid corals in the San Blas Islands,Panama

Mucous sheets are common on poritid corals as well as other reef coelenterates, but the frequency, cause and function of these sheets have rarely been investigated.Porites furcata Lamarck andPorites astreoides Lamarck colonies at three sites in the San Blas Islands, Panama, were monitored for the presence or absence of mucous sheets every 2 d between 11 May and 27 August 1983. Concurrent measurements of polyp expansion, particulate matter concentrations, salinity, water temperature and sedimentation rates were also made at each site. Mucous sheet formation was not correlated with any of the environmental parameters measured. Mucous sheet formation by both species had a lunar periodicity.P. furcata formed mucous sheets around the time of the full moon, whileP. astreoides formed mucous sheets during the first quarter (around a week before full moon). Mucous sheet formation was correlated with polyp contraction behavior and may be a secondary effect of a lunar cycle in polyp activity.     

Night irradiance and synchronization of lunar release of planula larvae in the reef coral Pocillopora damicornis

Pocillopora damicornis (Linnaeus), which is known to release planula larvae on a monthly cycle, was grown in full daytime solar irradiance, but with four treatments of night irradiance: (1) natural night irradiance, (2) shifted-phase (total darkness during nights of full moon with artificial irradiance at lunar intensity on nights of new moon), (3) constant full moon (full lunar irradiance every night), and (4) constant new moon (total darkness every night). The reproductive cycle of the corals held in the shifted-phase treatment moved out of synchrony with the cycle of corals exposed to a natural lunar cycle of night irradiance. Two previously described types of P. damicornis were tested. The Type Y normally start releasing larvae at full moon, with peak production at third quarter. In the shifted-phase treatment they began releasing planulae at new moon (artificial full moon), with peak production at first quarter. The Type B corals, that normally start releasing planulae at new moon with peak production at first quarter, began to release planulae at full moon (artificial new moon), with peak production at third quarter. Populations of corals grown either in the constant full moon or constant new moon treatment quickly lost synchronization of monthly larva production, although production of planulae continued. Thus spawning is synchronized by night irradiance.Contribution No. 702 of the Hawaii Institute of Marine Biology     

Sublethal predation on Polydora cornuta (Polychaeta: Spionidae): patterns of tissue loss in a field population,predator functional response and potential demographic impacts

Patterns of tissue loss due to sublethal predation and potential effects on population dynamics were investigated in the polychaete Polydora cornuta Bosc, 1802. Field observations in southeastern Connecticut showed that the percentage of adult worms regenerating segments and/or feeding palps varied temporally from May to November 1982, but the general pattern suggests a constant level of sublethal encounters. Over all sampling dates, a mean of 14.9 and 7.0% of the population was found regenerating posterior segments and palps, respectively. Worms lost a mean of 19.1% of their segments. There was a weak, but statistically non-significant, size-specific difference in the number of regenerating segments per individual. Worms with 20 to 39 segments and 60 to 80 + segments lost an average of 13.8 and 17.2% of their segments, respectively, but worms with 40 to 59 segments lost an average of 23.4% of their segments. The incidence of regeneration (palps and segments) increased linearly with increasing adult density in the population. There was also a strong positive linear relationship between the number of segments available and the number of segments lost at the population level. This suggests that the constant level of sublethal predation can be explained by predators taking prey in proportion to their density and/or the number of segments available. This pattern is supported by results from a laboratory functional response experiment which indicated that at high prey density, partial consumption of P. cornuta by the predatory polychaete Eteone heteropoda was greater than lethal consumption. Lethal predation of P. cornuta by E. heteropoda did not vary across experimental density treatments. Demographic modelling suggested that sublethal predation can reduce the population growth rate of P. cornuta, but the reduction is less than if the added predation pressure was solely lethal. It was estimated that up to 25% of the population could be preyed upon in a sublethal manner before the potential for population growth fell below population maintenance levels.     

Temporal patterns and transport processes in recruitment of spiny lobster (Panulirus argus) postlarvae to south Florida

We used time-series analysis to identify weekly and annual patterns in the supply of spiny lobster, Panulirus argus (Latreille, 1804), postlarvae to the Florida Keys, USA, over an 8 yr period. We also investigated the relationship between postlarval influx and wind forcing as a transport mechanism using the complex vector-scalar correlation analysis. Postlarval supply had a lunar phase periodicity at 4.5 wk intervals, with postlarval abundance peaking between the new moon and first-quarter lunar phases. A distinct annual cycle of postlarval supply with two peak periods was also apparent. Cross-correlation analysis between relative postlarval abundance and a 12 mo cycle showed that the annual peak occurs in spring, centered around March. With the 12 mo periodicity removed, a smaller peak at 5 mo intervals was also well defined. Wind-forcing for 7 d prior to the time of postlarval collection was marginally correlated with postlarval abundance through the entire time-series; the association was strongest during the late fall to early spring months. The analysis indicated that postlarval supply was correlated with winds from the northeast (ca. 45°), which are associated with winter atmospheric fronts. In contrast to results reported for other spiny lobster populations, these patterns suggest that recruitment of lobster postlarvae to south Florida is predictable only at a gross level and is presumably affected by the temporally inconsistent structure of regional oceanic gyres and variability in the timing of lobster spawning in the Caribbean. Received: 24 January 1997 / Accepted: 4 March 1997     

Feeding responses of the tentaculate deposit-feederEupolymnia nebulosa (Annelida: Polychaeta): Influence of sexual maturity

Feeding responses of the tentaculate depositfeeding polychaeteEupolymnia nebulosa (Montagu) were studied by measuring rates of uptake of three different¹⁴C-labelled diatoms (unialgal cultures ofNavicula incerta Grunow,Nitzschia acicularis Wm Smith, andNitzschia sp.). Worms used during this study were collected in the harbor of Port-Vendre (Western Mediterranean) during August 1986 (immature worms) and December 1987 (mature worms). Uptake rates were affected both by the length of the experiments and by the nature of the food offered. The highest rate of uptake (2.98 10^–4 mg ashfree dry wt of algae mg^–1 dry wt of worms h^–1) was obtained during short-term experiments (4 h) with the smallest diatom (Nitzschia sp.). The lowest rate of uptake (0.21 10^–4 mg ash-free dry wt of algae mg^–1 dry wt of worms h^–1) was also obtained withNitzschia sp., but for a long-term (48 h) experiment. There was no significant difference between rates of uptake of immature and mature worms.     

Induction of swarming ofNereis succinea

Nereis succinea (Frey and Leuckart, 1847), collected in 1987 from the Weser estuary, FRG, was exposed to different temperatures in the laboratory. Metamorphosis to heteronereid stages, as well as swarming at a minimum temperature of 12°C, was induced by raising temperatures around the time of the new moon. Lunar periodicity was illustrated under natural temperature-programs, and at 16°C. An abrupt increase in temperature caused swarming to occur at different times of the lunar cycle.     

Anterior regeneration in the spionid polychaetes<Emphasis Type="Italic"> Dipolydora quadrilobata</Emphasis> and<Emphasis Type="Italic"> Pygospio elegans</Emphasis>

Like many other benthic infaunal invertebrates, spionid polychaetes often lose portions of their bodies to predators, which affects their activities. Feeding palp loss and tail loss have been studied in several spionids, but the capacity for anterior tissue regeneration has not been compared in different species. The present study examines anterior tissue regeneration in two species, Dipolydora quadrilobata (Jacobi 1883) and Pygospio elegans Claparède 1863, in two laboratory experiments. Tissue removal treatments included removal of palps only, removal of anterior tissue through the first setiger, anterior tissue through the fifth setiger, all anterior tissue through half of the gill-bearing setigers, and all anterior tissue through the last gill-bearing setiger. Regeneration was monitored by capturing images of the worms and digitizing the area of regenerated anterior tissue or counting the number of segments that grew over time. Worms of both species regenerated anterior tissue regardless of the amount removed. Morphogenesis during regeneration followed a similar pattern in these two species regardless of the amount of anterior tissue lost, progressing from wound healing to formation of a recognizable prostomium and peristomium (“head”) by 6 days post-ablation. Palp and setal growth, addition of segments, and formation of nuchal organs and the ciliated food groove followed so that worms appeared to have re-grown “normal,” but smaller, “heads” and palps by 9–12 days following ablation. Over the course of 16 days, worms that lost more segments regenerated less tissue relative to their initial intact size and did so more slowly. There was no significant palp growth during the first 3 days following ablation. Rate of segment addition was directly related to the degree of tissue loss in D. quadrilobata. P. elegans added segments at similar rates whether 50 or 70% of the original segments was removed.     

Migration of the sandworm Nereis virens during winter nights

There have been many previous reports of the sandworm Nereis virens Sars swimming in the water column. This behavior usually has been attributed to reproductive processes. Sandworms were found swimming in surface waters at night on ebb tides during many nights of January, February and March in a Maine (USA) estuary. None of the specimens examined contained gametes or possessed other characteristic spawning or pre-spawning modifications. Several age classes were found, with worms measuring 9 to 38 cm in length, weighing 0.5 to 19.8 g, and having 82 to 187 segments. The greatest numbers of worms were observed during near-average tides on evenings in which low tides occurred a few hours after sunset but prior to moonrise. Up to 83 worms per minute were observed swimming seaward through a 20 m transect, while none were observed swimming landward at any stage of the tide. It is concluded that sandworms swimming during winter nights is unrelated to reproduction and that it is an inherent behavior pattern.Contribution No. 102 of the Ira C. Darling Center, Walpole, Maine 04573, USA.     

Trace-metal detoxification and tolerance of the estuarine worm<Emphasis Type="Italic"> Hediste diversicolor</Emphasis> chronically exposed in their environment

Polychaete worms (Hediste diversicolor) originating from a strongly metal-contaminated area (Restronguet Creek) and a relatively clean site (Blackwater estuary) were exposed to a range of experimental doses of Ag, Cd, Cu and Zn. Specimens from both populations were compared to assess their relative sensitivity to metal stress and the physiological mechanisms involved in their respective adaptive strategies for coping with increased metal exposures. Taking into account the LC₅₀ values, increased tolerance to Cd, Cu and Zn of the Restronguet Creek worms over that of the Blackwater was demonstrated, whereas the opposite was shown for Ag. An abundant secretion of mucus in response to toxicants was observed, possibly reducing metal availability for uptake, at least under laboratory conditions. This mechanism was particularly active in specimens from Restronguet Creek exposed to Ag and Cu. Unexpectedly, of the two worm populations, Blackwater worms contained significantly higher concentrations of cytosolic heat-stable compounds (CHSTC), a category of cytosolic components that includes metallothioneins, the detoxificatory role of which is well documented, and other compounds binding trace metals via sulphur bonds. However, the concentration of such compounds is not totally representative in itself of their involvement in metal detoxification, because a high rate of their turnover in metal-exposed worms might be responsible for (at least) the Cu storage associated with S in lysosomes as a consequence of the breakdown of Cu-thionein. Cu-containing lysosomes were abundant in epidermal cells of Restronguet Creek worms, but were lacking in Blackwater worms. Extracellular granules present in the epicuticle also contained S and Cu, their number and size being much more important in Restronguet Creek worms, and they appear to be a major detoxificatory store for accumulated Cu. Spherocrystals in cells of the gut wall seem to be the major detoxified store of Zn in Restronguet Creek worms, whereas in specimens from the Blackwater they were also present but contained only S and Ca at detectable levels. Evidence was also found for the presence of detoxificatory intracellular structures containing other metals and metalloids in the tegument and gut epithelium of Restronguet Creek worms.Communicated by S.A. Poulet, Roscoff     

Chemical defense and evolution in the Sacoglossa (Mollusca: Gastropoda: Opisthobranchia)

Summary. Recent advances in both the systematics and the natural products chemistry of the order Sacoglossa (=Ascoglossa) in the gastropod subclass Opisthobranchia suggest a revised and improved historical account of the evolution of the group. Although the algal genus Caulerpa makes a suitable model for the ancestral food of the order, other siphonaceous algae are consistent with both morphological and chemical data. At an early evolutionary stage terpenoids are sequestered from the food, and used defensively, often with modification. With an evolutionary switch to different kinds of algal food, there is often a shift to other, related defensive chemicals. A switch to new food source sometimes leads to the abandonment of chemical defense, but in other cases there is de-novo synthesis of defensive metabolites. The synthesis of polypropionates, which are used defensively, occurs in some other gastropods, but otherwise is known only in fungi. The systematic distribution of the defensive polypropionates suggests that their defensive use has evolved several times among gastropods. Failure to detect them may mean that synthetic capacity has evolved more than once, or it may mean that they exist at low levels, perhaps having a non-defensive function. Received 9 February 1998; accepted 20 March 1998.     

Population ecology and secondary production of the polychaete Loimia medusa (Terebellidae)

In soft-sediment marine and estuarine habitats, population dynamics of resident species are an important aspect of commnity structure and function, yet the population dynamics of many members of these communities remain poorly studied. The population dynamics and secondary production of the infaunal terebellid polychaete Loimia medusa (Savigny) were investigated in a shallow sand habitat of the York River, Chesapeake Bay, Virginia, between June 1989 and December 1990. Monthly sampling throughout 1 yr revealed that individuals were present year-round with maximum densities of 60 ind m^-2 and a life span of 1 yr. Abundances were highest from August through September when two cohorts recruited to the population. Mean individual growth was highest in summer, and appeared to be a function of temperature rather than food limitation. Adults were reproductive from May through October and exhibited high fecundity; the number of oocytes segment^-1 ranged from 201 to 15840 among worms. Larvae initially appeared approximately 1 mo after gametes were first observed in adult L. medusa, and these larvae appeared to spend less than 1 mo in the plankton. Abundances of larvae in the water column were significantly higher around new moon, suggesting lunar spawning periodicity. New recruits attained sexual maturity within 2 mo of settlement. Secondary production was 3.3 g ash-free dry wt m^-2 yr^-1 and the P/B (production/biomass) ratio was 3.0. Despite equilibrium species characteristics such as large size, high fecundity, and planktonic larvae, L. medusa exhibits some characteristics typically ascribed to opportunistic species, namely, the capacity for rapid growth and maturation, a short life span and a relatively high P/B ratio. This mixture of life history characteristics emphasizes that there is no simple dichotomy between opportunistic and equilibrium species. We caution that the dominance of large infaunal polychaetes in soft-bottom estuarine and marine environments may not be a useful indicator of undisturbed habitats.     

Gametogenic periodicity in the chemosynthetic cold-seep mussel “<Emphasis Type="Italic">Bathymodiolus” childressi</Emphasis>

The gametogenic periodicity of the cold seep mussel “Bathymodiolus” childressi was analysed from a time series of samples from depths of ∼ 650 m surrounding the Brine Pool cold seep on the continental slope of the northern Gulf of Mexico. Occasional samples were retrieved from Bush Hill and GC 234 for comparison. At the Brine Pool, both females and males showed strong reproductive periodicity with the initiation of gametogenesis from December to March, followed by a period of gamete growth or proliferation and spawning from October to February. Gametogenesis was synchronized at all three sites. Gametogenic periodicity appears to be correlated with surface production that peaks during the winter months. Downward flux of detritus during this period may provide food for the planktotrophic larvae and also supplementary nutrition for the adult, which has chemosynthetic bacterial symbionts but is also capable of filter feeding. Individuals in all three populations carried parasites and these were especially common at Bush Hill and GC234, where it is suggested they have a major impact on reproductive output.     

Gametogenesis and spawning periodicity in the fan worm <Emphasis Type="Italic">Sabellastarte spectabilis</Emphasis> (Polychaeta: Sabellidae)

The sabellid polychaete Sabellastarte spectabilis (Grube 1878) was collected at approximately monthly intervals from January 2002 to December 2003 from intertidal and subtidal reefs near the Hawaii Institute of Marine Biology in Kane’ohe Bay, Hawaii, USA (21°N, 157°W). Gametogenesis and spawning periodicity were investigated using histological techniques and induction of spawning trials. Worms were characterized into four discrete reproductive stages based on histological evidence: (1) No evidence of reproductive activity in the coelom (sex cannot be determined), (2) Only coelomocytes present in the coelom (sex cannot be determined), (3) Some gametes present in the coelom (sex can be determined) and (4) Coelom densely packed with gametes (sex can be determined). The small hermaphroditic portion of the population was not used in this study. Stage 4 worms were present over an extended period of time (females, March–December and males, March–November) indicating a potentially broad reproductive season. No correlation between day length and maturation stages in S. spectabilis was detected. However, the statistical model Y = ([394.26 × X] − [7.793 × X ²]) − 4960.781 where Y the % frequency of Stage 4 worms and X the mean monthly water temperature explained 44% of the variation between water temperature and % frequency of Stage 4 worms. Maturation appeared to coincide with water temperatures of 24–25°C (March–September) after which there is a reduction in the % frequency of stage 4 individuals. Induction of spawning trials conducted between May and January showed the month of October with a significantly higher percent success than any other month investigated. According to all available information (e.g., natural spawning in water tables, histological data, induction of spawning trials, correlation of maturation stages with observed changes in average monthly water temperature.), there is an apparent peak in reproductive activity (spawning) within a broad maturational season, which may be influenced by water temperature.