Sexual reproduction,larval development and release in <Emphasis Type="Italic">Spongia officinalis</Emphasis> L. (Porifera,Demospongiae) from the Apulian coast

The reproductive cycle of the bath sponge Spongia officinalis L. has been studied over 1 year on 11 tagged specimens of different sizes (from 82 to 886 ml, in volume) from Ionian coasts of Apulia (SE Italy). According to literature data, the sponge is viviparous. All the monitored specimens showed sexual reproduction, even if the process usually involved small portions of the sponge tissue. Ten specimens were gonochoric (sex ratio 1:1), one specimen showed successive hermaphroditism, with alternate production of oocytes and spermatic cysts in the same reproductive season. Young oocytes occur almost all year round, whereas large mature eggs show a peak in October–November, concomitantly with the appearance of spermatic cysts. No relationships were observed between the sponge size and the presence of sexual elements within the range of the sponge size considered in this research. Embryo development occurs in patches of choanosomal tissue, which contain four or more elements. Cleavage is total and equal; it starts in November and in May leads to a solid stereoblastula, which develops into a parenchymella larva, from May to July. The stereoblastula lacks flagella and its surface is delimited by elongated cells well segregated from the internal ones. Parenchymella larvae are released from June to July, asynchronously, either at the individual or population level, with a few days of de-phasing. Up to 523 larval elements/48 h for a sponge specimen were counted. The free-swimming larvae are ovoid and uniformly flagellated. Flagella are longer at the posterior region, than on the rest of the larval body. Flagellated cells form a pseudo-stratified epithelial layer, delimiting the outermost larval surface, and are filled with electron-lucent vesicles showing a homogenous content. No choanocyte chambers, pinacocytes or skeletal elements are present in the newly released larvae.     

Silica content and spicule size variations in Pellina semitubulosa (Porifera: Demospongiae)

 The variations in both silica content and spicular size were studied in two populations of the demosponge Pellina semitubulosa (Lieberkühn). Samples were collected over a period of 1 year (June 1994 to May 1995) in two Mediterranean coastal basins: Porto Cesareo (southwestern Apulia) and Marsala (northwestern Sicily). The values of spicule size (length and width) and sponge silica content were significantly higher in the population of Porto Cesareo, where the highest water silica concentration was recorded. In both Porto Cesareo and Marsala the sponge silica content showed a seasonal trend, positively correlated with water temperature values. In both populations, the smallest spicules were found in specimens collected from summer to late autumn, after sexual reproduction. Secretion of new spicules may be connected with the process of remodelling occurring in sponges after gamete and larval release. Received: 10 October 1999 / Accepted: 13 April 2000     

Reproductive strategies of Mycalecontarenii (Porifera: Demospongiae)

The sexual and asexual phases of the reproductive cycle of the sponge Mycalecontarenii (Martens, 1824) were studied from samples collected over a period of 2 years (June 1994 to May 1996) in a Mediterranean coastal basin (Porto Cesareo, southwestern Apulia). The species is viviparous; year round it shows a discontinuous oocyte production. No males were found, possibly owing to the very short period of spermatogenesis. M.␣contarenii produces asexual buds during autumn and winter, and this process seems to be related to changes in water temperature. Even though, at present, we cannot evaluate the incidence of larval recruitment, we postulate that asexual reproduction in M.contarenii plays a crucial role in maintaining the sponge population. Received: 8 July 1997 / Accepted: 5 November 1997     

Predictable annual mass release of gametes by the coral reef sponge Neofibularia nolitangere (Porifera: Demospongiae)

Mass release of gametes of the sponge Neofibularia nolitangere (Duch. & Mich., 1864) occurs simultaneosly along the leeward coast of Curaçao over a period of three subsequent days, without any apparent spatial pattern in the sperad of activity. A population of 99 individual sponges was monitored from August through November 1984 for development and subsequent release of gametes. Release started every day of 3 3-d period (12 to 14 October) at about 1400 hrs and lasted until just after sunset (1830 hrs). Ninety percent of the population showed reproductive activity. Exactly one lunar month later (11 to 12 November), a second release of gametes occurred. In the following year the same sequence of events was observed for the original population (2 to 4 October and 1 to 2 November, 1985). In all instances the first gamete release began on the third day after the full moon. These and earlier observations on this phenomenon show a strong correlation between moon phase and the time of gamete release. Histological and field observations show separate development and release of gametes (sex-ratio males: females 1.5:1) with external fertilization. No sex-reversal occurred in the population during release activity the following year, demonstrating a clear gonochoristic and oviparous type of reproduction for N. nolitangere. First development of oocytes and spermatocytes has been observed at 36 and 7 d, respectively, before the date of first release. In both males and females, a major part of the mesohyl of the total sponge was involved in gamete development. Spermatozoa are released through the osculum as a dense white smoke; eggs consisting of oocyte-nurse cell globules, reinforced with spicules, are separately released en masse with the outgoing water stream of the female sponge. Shortly after release the eggs become sticky and show a negative buoyancy. The reproductive strategy of N. nolitangere appears to be directed at maximizing the number of surviving recruits by maintaining a high reproductive output at a short specific time interval.     

Variability in reproductive output across a water quality gradient for a tropical marine sponge

To establish a complete understanding of reproductive variability, larval supply and ultimately population demographics of a species it is important to determine reproduction across a broad spectrum of environmental conditions. This study quantified sexual reproduction of the brooding, gonochoristic sponge Rhopaloeides odorabile from populations across the shelf reefs of the central Great Barrier Reef (GBR). Histological sections of reproductive sponges collected at increasing distances from the coast were used to determine if numbers of reproductive sponges, reproductive output (using a reproductive output index), size at sexual maturity, and sex ratios varied according to their location (distance) from the coastline and therefore from influences of terrigenous/riverine discharge. Significantly higher proportions of reproductive sponges occurred with increasing distance from the coast. The proportion of all reproductive sponges (both male and female) on offshore reefs ranged from 77 to 90%, during November and December, the peak reproductive months of this sponge, compared to 47 to 50% for sponges occurring on coastal reefs. Levels of female reproduction increased with increasing distance from the coastline on two levels. First, oocytes from offshore sponges were significantly larger than oocytes from coastal sponges. Second, sponges from offshore reefs showed a reproductive index (proportions of oocytes, embryos and larvae mm⁻²) approximately 15 times higher than coastal reef sponges. Therefore, both numbers of oocytes, embryos and larvae in conjunction with larger oocytes contribute to a higher reproductive output index for offshore sponges. The production of spermatic cysts in males was consistent across the GBR. Sex ratios for coastal reef sponges showed a male bias while offshore sponges showed approximate equal sex ratios. The effect of terrigenous riverine input from coastal fluvial plains to the inner GBR is well established and is likely to contribute to the lower levels of reproduction associated with female sponges inhabiting coastal reefs of the central GBR.     

Effect of diet on growth rate and reproductive fitness of Turbo sarmaticus (Mollusca: Vetigastropoda: Turbinidae)

Experiments were conducted to determine the effect of four algal diets (Corallina spp., Gelidium pristoides, Ulva rigida and an equal volume mixture of these seaweeds) on growth and reproduction of the South African gastropod Turbo sarmaticus Linnaeus, 1758. The best growth rate of juveniles (up to 13.8 mm shell length and 34.26 g wet body weight increase in 12 months), reproductive fitness of mature specimens (gonad index up to 33%) and energy reserves (up to 4.76 mg glycogen/100 mg foot tissue) were achieved when T. sarmaticus was fed U. rigida or a mixed diet. In addition, the gonad index of individuals fed these diets was almost twice that of similar-sized field specimens. Juveniles fed Corallina spp. only, grew very little (only 2.4 mm shell length and 4.23 g wet body weight increase in 12 months). The reproductive fitness of adults fed on such a diet was also poor (gonad index <4.5%) and energy reserves were low (<3.5 mg glycogen/100 mg foot tissue). Along the southeastern coast of South Africa, T. sarmaticus has a distinct reproductive cycle with gametogenesis occurring from March/April until August/September, whilst maturity (gonad index = 15%) was maintained until December, after which spawning occurred until March. Received: 6 July 1998 / Accepted: 8 March 1999     

Larval reproduction in the ctenophore Mnemiopsis mccradyi (order Lobata)

Dissogony was observed in normal and experimentally manipulated individuals of Mnemiopsis mccradyi in February–April, 1985. The normal oral-aboral length at which sexual reproduction begins in M. mccradyi is 30±5 mm; however, some larvae, of 1.5 to 2.8 mm (oralaboral length), produce viable gametes. During the present study, not all individuals from a particular spawning which were kept in the same conditions became reproductive at an early age. The age at which larval reproduction began varied between individuals. The length of time over which individuals spawned varied considerably — some spawned only once, while others spawned repeatedly. Only the gonads located under the adesophageal ctene rows (and not those associated with the adtentacular rows) produced gametes during the early reproductive period. After larvae stopped spawning, the gonads regressed and the larvae grew to large lobate stages. Individuals which became sexually reproductive as larvae also became sexually reproductive during lobate stages.     

Reproduction and population dynamics of the fissiparous brittle star Ophiactis savignyi (Echinodermata: Ophiuroidea)

Reproductive cycle, asexual reproduction, and population dynamics of the fissiparous brittle star, Ophiactis savignyi, which inhabits the exhalant passages of the sponge Haliclona sp. were examined monthly from February 1991 to January 1992 at Wanlitung, southern Taiwan (22°N; 120°E). Mature gametes were found from March to December, but release was mainly in May and June. Sexual recruits were found from May to December, with the highest frequency (14.1% of population) in June. Sexual recruits composed 2.4% of the 1-yr sample. Sex ratio of male to female was 24:1. Fission occurred throughout the year, although the frequency of recently split individuals was lower from January to June (6 to 31%), and higher from July to December (42 to 52%). The occurrence of fission was highest after spawning. Regenerating individuals composed 48.2% of the 1-yr sample. Population density fluctuated greatly during summer due to recruitment by fission and mortality or dispersal due to the stressful environmental conditions. Both sexual and asexual reproduction of O. savignyi were successful at this site.     

Sessile and non-sessile morphs of Geodia cydonium (Jameson) (Porifera, Demospongiae) in two semi-enclosed Mediterranean bays

Morphological plasticity and ecological aspects of the demosponge Geodia cydonium (Jameson) were studied from seasonal samples collected over 1 year in two semi-enclosed Mediterranean bays of the Southern Italian coast (Marsala lagoon and Porto Cesareo basin). Sponge specimens present two morphs: sessile and non-sessile, both of which showed constant size distribution and density over the studied year. Sessile specimens were larger in size than non-sessile ones. This feature is particularly evident at Porto Cesareo, where these sponges have a more compact skeletal network than at Marsala (evident both in the cortical spicule size and sponge silica content). Sessile specimens adhere to hard rocky substrates (Porto Cesareo) or phanerogam rhizomes (Marsala); non-sessile ones occur on soft bottom areas. Several morphological and structural features of the non-sessile forms differ in the two environments, but the difference in body shape seems to play the most relevant role in enhancing the colonization of incoherent substrates. Indeed, at Marsala, where the large amount of silt and clay determines the occurrence of a markedly reduced anoxic layer just below the surface of the sediment, non-sessile specimens of G. cydonium are fairly spherical and thus able to roll, dragged by slow circular currents. In addition, the usual association with the red alga Rytyphlöea tinctoria, which almost constantly forms a thick and continuous layer around the sponge, allows them to avoid contact with the substrate. The non-sessile specimens from Porto Cesareo inhabit sandy soft bottoms and are flattened. In such an environment, affected by moderate wave turbulence, the flattened shape widens the contact surface between the body and the substrate, thereby reducing the risk of stranding. The evident signs of abrasion, provided by scanning electron microscopy investigations, on both cortical spicules and outermost sponge surface suggest that sponges rub on the bottom. Sediment, epibiontic organisms, and the phanerogam leaves protect this sciaphilous sponge from high solar radiation, allowing the specimens to live in these shallow environments.     

Co-occurrence of three types of egg policing in the Norwegian wasp <Emphasis Type="Italic">Dolichovespula norwegica</Emphasis>

In insect societies, workers often try to challenge the reproductive monopoly of the queen by laying their own eggs. Successful worker reproduction, however, is frequently prevented by queen policing or worker policing, whereby either the mother queen or non-reproductive workers selectively kill worker-laid eggs. Recently, a third mechanism—“selfish” worker policing—has also been described in which the workers selectively police worker-laid eggs but also lay eggs themselves. Here, we present results from the monogynous wasp Dolichovespula norwegica, which show that all three kinds of policing—queen policing, worker policing and “selfish” worker policing—co-occur. The net effect of these three kinds of policing collectively favoured the queen’s reproduction, as within 1 day 44% of the worker-laid eggs versus only 8% of the queen-laid eggs were eaten. Of the worker-laid eggs that were killed by workers, approximately two thirds were eaten by the reproductive workers even though these made up only a small proportion, 8%, of the work force. This means that policing workers obtained both direct fitness benefits as well as indirect (inclusive) fitness. In addition, we show that worker policing was carried out by a limited, specialised set of workers that was estimated to constitute approximately one quarter of the whole colony and of which 66% were non-reproductive.     

Reproduction and growth of Isostichopus fuscus (Echinodermata: Holothuroidea) in the southern Gulf of California, México

The reproduction and growth of Isostichopus fuscus (Ludwig, 1875) at Isla Espíritu Santo, Baja California Sur, México from July 1992 to September 1993 and from August 1996 to July 1997 were analyzed. The reproductive cycle was monitored by using histological analysis. Five gonadal stages were described according to cell types present in the gonad; undetermined, gametogenesis, maturity, spawning and post-spawning. This holothurian reproduces annually during summer (July to September), when sea surface temperature reaches 27 °C. The gonad index is related to the gonadal stages. The first sexual maturity for I. fuscus was calculated at 367 g (4 to 5 years old) according to the von Bertalanffy equation, and nearly all the specimens at Isla Espíritu Santo were 5 to 7 years old (including some individuals over 35 years old). Data of the length–weight relationship showed that I. fuscus grows allometrically at the studied site (b = 1.83). The mortality rate was relatively high (M = 0.51), indicating that in a typical cohort, 40% of the recruits does not survive the first year, and less than 20% reaches maturity. Received: 15 July 1998 / Accepted: 19 August 1999     

Sexual and asexual reproduction of<Emphasis Type="Italic"> Allostichaster capensis</Emphasis> (Echinodermata: Asteroidea) in Golfo Nuevo

Sexual and asexual reproduction of the fissiparous starfish Allostichaster capensis were examined for 24 months at Bahía Kaiser, Golfo Nuevo, Chubut, Argentina. Gonad indices revealed an annual reproductive cycle, with peak development occurring in late winter (August). Spawning occurred in spring (September). Pyloric caeca indices displayed a clear annual cycle and a reciprocal relationship with gonad indices, showing the storage function of pyloric caeca. The starfish population appears to be largely maintained by fission, as only two females were found. A. capensis also showed an annual cycle of fission, with the highest frequency in spring and summer (from November to January), just after spawning. Incidence of fission was correlated with environmental factors such as photoperiod, seawater temperature, and salinity.Communicated by P.W. Sammarco, Chauvin     

Sexual reproduction of the solitary sunset cup coral Leptopsammia pruvoti (Scleractinia, Dendrophylliidae) in the Mediterranean. 2. Quantitative aspects of the annual reproductive cycle

There are few data on the reproductive biology of coral species living in temperate zones, particularly in the Mediterranean Sea. Leptopsammia pruvoti is a solitary coral that is commonly found in sea caves and under overhangs throughout the Mediterranean basin and along European coasts from Portugal to southern England. In this paper, we describe its annual reproductive cycle in the eastern Ligurian Sea near the city of Leghorn (Tuscany, Italy). Polyps were sexually mature at 3 mm in length (maximum diameter of the oral disc), were dioecious with a sex ratio of 1:1, and brooded their larvae. The maturation of spermaries took 12 months and oocytes 24 months. The rate of gonad development increased significantly from November to January, fertilization occurred from January to April and planulation during May and June. Seasonal variations in water temperature and photoperiod may have played an important role in regulating reproductive events. The amount of energy devoted to male gametogenesis (quantified by gonad index) was significantly higher in the sex separated species L. pruvoti than in the hermaphroditic dendrophylliid Balanophyllia europaea, whose reproduction has been studied in a previous work. We hypothesize that this difference is due to the contrasting sexuality or fertilization biology of these two species (cross-fertilization in the sex separated L. pruvoti versus possibly self-fertilization in the hermaphroditic B. europaea). Greater male sexual allocation in reproductive strategies characterized by dioecism or cross-fertilization when compared to those characterized by hermaphroditism or self-fertilization is common in plant mating systems. In relation to other solitary dendrophylliids, L. pruvoti presents an ‘r’-reproductive strategy.     

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.     

Seasonal variation in the effects of food availability on gametogenesis in the purple urchin (<Emphasis Type="Italic">Strongylocentrotus purpuratus</Emphasis>)

The amount of food and when it is available affect both the timing of reproduction and the number of gametes produced by purple sea urchins, Strongylocentrotus purpuratus. To investigate this further, the effects of food availability on feeding rates, gonad growth, and gamete development were examined in S. purpuratus collected from the Point Loma kelp forest near San Diego, California, USA (32.69° N, 117.26° W) in September (Fall) 2007, and February (Spring) and July (Summer) 2008, using laboratory mesocosms. Each seasonal laboratory feeding experiment lasted 3 months, and different levels of food availability were established with different feeding frequencies (from 1 to 7 days week⁻¹). Gonad tissues of male and female urchins were staged at the end of each experiment using histological analyses. Reduced food availability resulted in increased daily consumption rates, especially in the Fall when gamete development began. Food limitation at this time resulted in failure to produce viable gametes, suggesting there is a critical period early in gonad development when food limitation affects reproductive competency. Food limitation later in gonad development did not stop viable gamete production, although it did reduce gamete output.     

Multiple spawning events and sexual reproduction in the octocoral Sarcophyton elegans (Cnidaria: Alcyonacea) on Lizard Island, Great Barrier Reef

Sarcophyton elegans is a common symbiotic (zooxanthellate) octocoral species in the shallow waters of the Great Barrier Reef (GBR). Study of a population at Lizard Island (14°40′S, 145°28′E) on the GBR from October 1991 to January 1994 revealed that, as is typical of tropical alcyonarian corals, S. elegans is a gonochoric broadcast spawner with a 1:1 sex ratio. Sexual reproduction was closely correlated with colony size, with first reproduction at 13-cm basal stalk circumference for females and 12 cm for males. Oogenesis took 19–24 months, with a new cycle commencing every year, and spermatogenesis took 10–12 months. The majority of gametes were released during the annual austral mass coral spawning event after the full moon in November, but gametes were also released after the full moon in each month between August and February. All autozooid polyps participated in reproduction, but those at the outer edge of a colony released their gametes first. During subsequent months, the polyps closer to the center of the colony released their gametes. This is a novel strategy of gamete release, reported here for the first time, which accommodates the demands of feeding and reproduction in a different way than other corals where individual polyps have separate feeding or reproductive roles. Colonies upstream in the prevailing current spawned up to 1 month earlier than those downstream and ceased 1 month earlier. The mechanism controlling this spatial differentiation in spawning time, repeatedly observed over three seasons, is unknown. Sarcophyton elegans appears to have a dual strategy of providing protection for its gametes by releasing most of them concurrently with the single, annual mass spawning of a large number of cnidarians, while also hedging its bets by individual colonies spawning a fraction of their gametes over an extended period of 6 months.     

Reproduction activity of<Emphasis Type="Italic"> Exogone naidina</Emphasis> (Polychaeta: Exogoninae) in a population from the Apulian coast (Adriatic Sea)

This study reports on the reproduction activity of Exogone naidina Örsted, 1845, followed on a hard-bottom population located at a site along the coast of Bari (South Adriatic Sea), during a year of study from November 2000 to October 2001. Population structure was analysed together with the reproductive state of individuals over time. The interpretation of frequency histograms computed on the total worm length was not easy because of the almost complete lack of juvenile forms. Ripe individuals were present for most of the year, covering a large range of sizes. The reproductive season extended from April to December, with two peaks of activity: one from April to June and one in September–October. Life span was hypothesized to be up to 1 year, with generational change clearly visible only in late summer. However, the prolonged reproductive period produces an overlapping of generations. The link between population density and the reproductive cycle of the species with food availability is also discussed.Communicated by R. Cattaneo-Vietti, Genova     

Reproductive investment reduces recuperation from exhaustive escape activity in the tropical scallop Euvola zizac

In scallops, mobilization of reserves from the adductor muscle to support maintenance and reproductive activity may impinge upon a major role of the adductor muscle, the movement of the valves during swimming and escape responses. The tropical scallop Euvola ziczac (Linnaeus 1758) invests energy reserves to different degrees during its two periods of reproduction each year. We evaluated the impact of reproductive investment on the escape responses (clapping capacity and recovery after exhaustive exercise) of E. ziczac sampled at different reproductive stages (immature, mature, spawned) during the two reproductive periods in 1997. We compared the escape response capacities with the levels of muscle energetic reserves (glycogen, proteins, and arginine phosphate) and muscle metabolic capacities [activity of the glycolytic enzymes: glycogen phosphorylase (GP), pyruvate kinase (PK), phosphofructokinase (PFK), octopine dehydrogenase (ODH), arginine kinase (AK), and the mitochondrial enzyme, citrate synthase (CS)]. Gonad size, gamete volume fraction, and levels of gonadal protein and lipid were greater for mature scallops during the first than during the second reproduction. Numbers of claps during escape responses (49–57) and levels of muscle arginine phosphate remained similar throughout the different reproductive stages in both reproductive periods. In contrast, recovery was slowed during gonadal maturation in both reproductive periods and during spawning in the first reproduction. Scallops generally took more time to regain their initial clapping capacity during the first (25–40 min) than during the second reproduction (20–30 min). Muscle glycogen decreased markedly during both gonadal maturation and spawning in both reproductions; whereas, muscle proteins decreased only in the first reproduction. The levels of most enzymes decreased during gonadal maturation in both reproductions, and also after spawning, particularly during the first reproduction. We concluded that gonadal maturation and spawning did not decrease clapping capacity of E. ziczac, but decreased its capacity to recover from exhaustive exercise most likely due to decreased levels of energetic reserves and a reduced metabolic capacity of the adductor muscle. Moreover, these effects were probably stronger during the first cycle because of the greater reproductive investment coincident with decreased food availability. Received: 28 April 2000 / Accepted: 21 September 2000     

Genetic differentiation among populations of a broadcast spawning soft coral, Sinularia flexibilis, on the Great Barrier Reef

The genetic structure of 12 reef populations of the soft coral Sinularia flexibilis (Octocorallia, Alcyoniidae) was studied along the Great Barrier Reef (GBR) at a maximum separation of 1,300 km to investigate the relative importance of sexual and asexual reproduction, genetic differentiation and gene flow among these populations. S. flexibilis is a widely distributed Indo-Pacific species and a gamete broadcaster that can form large aggregations of colonies on near-shore reefs of the GBR. Up to 60 individuals per reef were collected at a minimum sampling scale of 5 m at two sites per reef, from December 1998 to February 2000. Electrophoretic analyses of nine polymorphic allozymes indicated that genotypic frequencies in most populations and loci did not differ significantly from those expected from Hardy–Weinberg predictions. Analysis of multi-locus genotypes indicated a high number of unique genotypes (N _go) relative to the number of individuals sampled (N) in each reef population (range of 0.69–0.95). The maximum number of individuals likely to have been produced sexually (N*) was similar to the number of individuals sampled (i.e. N*:N ˜ 1), suggesting that even repeated genotypes may have been produced sexually. These results demonstrated a dominant role of sexual reproduction in these populations at the scale sampled. Significant genetic differentiation between some populations indicated that gene flow is restricted between some reefs (F _ST=0.026, 95% CI= 0.011 − 0.045) and even between sites within reefs (F _ST=0.041, 95% CI=0.027 − 0.055). Nevertheless, there was no relationship between geographic separation and genetic differentiation. Analyses comparing groups of populations showed no significant differentiation on a north-south gradient in the GBR. The pattern in the number of significant differences in gene frequencies in pairwise population comparisons, however, suggested that gene flow may be more restricted among inner-shelf reef populations near to the coast than among mid/outer-shelf populations further from the coast. Received: 10 July 2000 / Accepted: 5 October 2000     

Seasonal reproduction in deep-sea sponges-triggered by vertical particle flux?

The examination of oogenesis and reproductive pattern in three deep-sea demosponge species from the foot of the Barents Sea continental slope (Norwegian Sea, 2300 m, 75°18N; 9°55E), Thenea abyssorum, Trichostemma sol and Tentorium semisuberites, suggests that sexual reproduction is triggered by the vertical flux of particulate organic carbon at least in one species. All three species examined reproduce sexually and proved to be oviparous and gonochoristic. A considerable number of asexually produced reproductive units was found in Thenea abyssorum, but owing to morphological details the origin of these buds is equivocal, and asexual reproduction is therefore judged to be of minor importance. Oogenesis in the three species can be divided into two developmental periods and does not differ greatly from processes known from shallow water species, except for the almost complete absence of nurse cells. For Thenea abyssorum a distinct seasonal reproductive cycle was found. The onset of yolk accumulation coincides precisely with a maximum in vertical particle flux as detected in sediment trap studies (von Bodungen et al. 1994). In addition, prior to yolk accumulation the sponge tissue appears almost empty, without storage units. Once yolk accumulation (i.e. the second growth period) has started, gamete development and release are completed within a few weeks. Thus sexual reproduction is an important reproductive mode in deep-sea sponges, which may be triggered by the pulsed input of POC to this food-limited environment, the energy-dependent process of yolk accumulation being the most sensitive process.