Gametogenesis and larval development in Sabella spallanzanii (Polychaeta: Sabellidae) from the Mediterranean Sea

Sabella spallanzanii (Gmelin) is one of the best known and widely distributed Mediterranean polychaetes, but available data on its general biology has been inferred mostly from populations recently introduced to Australia. In the present paper, data on gametogenesis and larval development of a Mediterranean population from the Gulf of Taranto (Ionian Sea, Italy) are reported. Histological and electron microscopical analysis of gametogenesis showed that oogenesis, a long process beginning soon after spawning in February, is of extraovarian type. By contrast spermatogenesis is a very fast process lasting from September to December, when the coelomic cavity is completely packed with mature gametes and almost devoid of coelomocytes. During the period of spawning, some experiments of in vitro fertilisation were performed, always without success, while some of the collected females naturally spawned fertilised eggs (100%). Particularly interesting was the discovery of a small percentage of eggs which appeared fertilised but unspawned within the coelomic cavity of female. Investigation of the S. spallanzanii life cycle, from the naturally spawned fertilised eggs to embryonic and juvenile stages, was performed by daily microscopical observations and by SEM. Despite the presence of large eggs giving rise to a lecithotrophic larva, a long pelagic phase was observed, the longest found among Sabellidae. The larval development pattern is also described. Settlement began after about 2 weeks; metamorphosis occurred 10 d after settlement, when mucus was secreted abundantly and an external tube was formed. The long larval pelagic period and the development pattern, suggesting a high potential for dispersal, support both the introduction and invasive behaviour of the species in Australian waters. Received: 28 May 1999 / Accepted: 10 December 1999     

Reproduction and larval development of Marenzelleria viridis (Polychaeta: Spionidae)

The pelagic larvae of the polychaete Marenzelleria viridis (Verrill, 1873) are newcomers among the meroplankton of the Baltic Sea, where they sometimes achieve an abundance of up to 21x10⁶ ind m^-3 near the coast, especially in late autumn and even in winter. Benthos samples were collected in the Darss-Zingst bodden chain from April 1992 to 1993 and used to ascertain the reproductive stages of individuals. Development from fertilized egg to benthic juvenile is described on the basis of field material. Details of the morphology of various pelagic stages and the young benthic worm are presented as drawings and scanning electron microscope photographs. In 1992, development of the gametes started in mid-May. The individuals reached maturity in late September after about 20 wk. The first planktonic stage was the fertilized egg. The larvae have initially one and later two pairs of black eyes and, in the 2nd segment, a ciliated pit. Gastrotrochs are present on the third and thereafter every alternate segment. Neuropodial setae develop once the 7-setiger stage is reached. Palps appear at the 10-setiger stage, and neuropodial bidentate hooks from the 10th to 11th setiger on. Metamorphosis into the juvenile benthic stage takes place at the earliest when the 15-setiger stage has been reached. The reproductive seasons of various populations of M. viridis and the developmental patterns of their larvae are discussed, and differences are compared with relevant findings for the spionids.     

Evolution of reproductive features and larval development in the genus Amphiglena (Polychaeta: Sabellidae)

Larval development in Amphiglena mediterranea is direct in the parental tube, and larvae crawl away as young juveniles with a radiolar crown capable of feeding. The non-feeding larvae of A. nathae are initially brooded in the parental tube, and then have a swimming phase before settling and developing a radiolar crown. Developmental data and reproductive features such as sperm structure are combined with other morphological data for an assessment of the cladistic relationships of the species in the genus Amphiglena. Two species of Laonome are included as part of the ingroup. Six genera, representing a sister group to the Laonome/Amphiglena clade, are used as an outgroup. Two most-parsimonious cladograms were found, and the evolution of reproductive features are discussed. The monophyly of Amphiglena is indicated by features previously identified, as well as two features based on sperm morphology. At present a plesiomorphic larval form cannot be identified for the genus Amphiglena. Received: 16 June 1997 / Accepted: 29 January 1998     

Effects of temperature on oocyte growth in the Mediterranean terebellid Eupolymnia nuebulosa (Annelida: Polychaeta)

Eupolymnia nebulosa (Montagu) is a widely distributed terebellid polychaete that builds its tubes on the coastal shelf in areas with mixed soft and hard bottoms. From a long-term survey in the Bay of Banyuls, France (NW Mediterranean), we found an advancement of the timing of the spawning period coincident with a delayed breakdown of the thermocline. We postulate that persistent high temperatures can influence gamete development by stimulating oocyte growth, resulting in earlier spawning. During 1992 and 1993, we used a between-individuals experimental approach to assess the possible effect of temperature on oocyte growth based on: (1) determination of the growing fraction of the oocyte population (i.e. oocyte net growth); (2) identification of differences in oocyte growth-rate among females; (3) comparison of non-significantly different size-distributions of the growing oocyte fraction (net oocyte size-distributions) at the beginning of the experiments with those at the end. No effect of temperature on oocyte growth was detectable at the population level, but a positive individual response to prolonged high temperature was evident. Thus, the lack of a significant response by the population to prolonged high temperature does not imply a lack of individual response. We propose a model of oocyte-growth dynamics based on temperature that incorporates previous observations of extended oogenesis and oocyte growth during periods of both increasing and decreasing temperature and on the scattered pattern of oocyte size-distributions at the onset of spawning. Received: 4 November 1996 / Accepted: 18 January 1997     

Reproductive isolation between intertidal and subtidal<Emphasis Type="Italic"> Scoloplos armiger</Emphasis> (Polychaeta,Orbiniidae) indicates sibling species in the North Sea

Two distinct modes of development in the common polychaete Scoloplos armiger (O. F. Müller, 1776) occur in the North Sea region: holobenthic development in egg cocoons and pelagic larvae hatching from suspended eggs. In the northern Wadden Sea near the island of Sylt, we observed that egg cocoons are produced intertidally while pelagic larvae originate from the adjacent subtidal zone. A previous genetic comparison between these subtidal and intertidal populations revealed distinct gene pools, suggesting that reproductive differences are not phenotypic but heritable. In this study, crossbreeding experiments show that intertidal and subtidal populations are reproductively isolated. Couples with males and females from different habitats had no offspring. Production of egg cocoons is determined by female origin from the intertidal zone. Pelagic larvae occurred only in couples with subtidal females and subtidal males. Intertidal males have spermatozoa with heads twice as long as those from subtidal males and a significantly shorter flagellum. We suspect that deviating sperm morphology may cause the reproductive breakdown at the fertilization stage. Juveniles hatching from cocoons have shorter anal cirri compared to juveniles that metamorphosed from pelagic larvae. We conclude there to be two sympatric sibling species in S. armiger: 'type I' in intertidal areas, which have egg cocoons, no pelagic larvae, elongated sperm heads, shortened sperm flagella and anal cirri; and a subtidal 'type S', lacking egg cocoons but with pelagic larvae, short sperm heads, long sperm flagella and anal cirri.Communicated by O. Kinne, Oldendorf/Luhe     

Genetic differentiation in Hediste diversicolor (Polychaeta: Nereididae) for the North Sea and the Baltic Sea

Genetic differentiation between North Sea and Baltic Sea Hediste diversicolor (O.F. Müller, 1776) (Polychaeta: Nereididae) populations was studied by allozyme electrophoresis on starch gel. Thirteen loci were analyzed in eight populations. The level of genetic variation was very low (mean H _o = 0.000 to 0.015). Differentiation between H. diversicolor populations is quite high (F _ST = 0.892) and reflected by three enzyme loci (MDH-I*, MDH-II*, IDH-I*). The reduced gene flow (N _m<1) may be explained by the limited dispersal capacity of the species. Regardless of whether found in the North Sea or Baltic Sea, there appear to be two different genetic types which are parapatric or sympatric in some places. The two types hybridize at three localities, but no signs of hybridization have been found at one (Tallinn). Received: 26 June 1997 / Accepted: 10 September 1997     

Microbial respiratory and ectoenzymatic activities in the Northern Adriatic Sea (Mediterranean Sea)

The carbon transfer through the microbial community in two areas of the Northern Adriatic Sea was estimated by proteolytic and respiratory activities during four oceanographic surveys carried out in June, 1996, 1997 and February, 1997, 1998. In front of the Po Delta (area A), the mean rates of proteolytic activity range from 4.9 to 9.9 r µg r C r h r l; near Ancona (area B), they range from 3.1 to 7.6 r µg r C r h r l. Respiratory rates vary between 0.19 and 2.29 and between 0.24 and 1.40 r µg r C r h r l in areas A and B, respectively. In general, high rates occur in the surface layers, within the first 10 r m of depth. In area A, proteolytic and respiratory rates undergo seasonal course, with high activity in warm periods. In area B, respiration and bacterioplankton abundance increase from the first to the second year, whilst proteolytic activity decreases. The sequence of metabolic steps in the carbon transfer within the bacteria, from the biotic vs . the abiotic compartment, was drawn in order to define the actual role of bacterial biomass in the biogeochemical fluxes in an ecosystem which often suffers distrophic crises. Respiratory turnover rates, in the upper 10 r m depth, reach low values in cold periods and high values in June, 1997. The carbon transfer versus mineralization flows better in the summer period, in particular in June, 1997. However, the bacterial growth efficiency ranges from 17 to 38% in area A and from 13 to 44% in area B with highest values in February, 1997, when bacteria contribute in a relevant way to the overall respiration.     

Larval growth of anchovy,Engraulis encrasicolus,in the Western Mediterranean Sea

Engraulis encrasicolus (Linnaeus, 1758) were sampled in July/August 1985 in the Western Mediterranean Sea; they were aged by means of growth rings in the sagittal otoliths. Daily growth rings were observed and subdaily rings were visible starting with the third or fourth daily increment. The Gompertz growth equation, commonly employed in larval growth analysis, suitably describes the growth of this species in the length range sampled; however, extrapolation to greater lengths is not reliable. An instantaneous growth rate of ca. 0.9 mm d^-1 was calculated for 8 mm larvae at a temperature of 20°C. This rate is higher than those reported by other authors for the related species E. mordax from colder waters.     

Oceanographic habitat,growth and mortality of larval anchovy (<Emphasis Type="Italic">Engraulis encrasicolus</Emphasis>) in the northern Aegean Sea (eastern Mediterranean)

Data from two ichthyoplankton surveys carried out during June 1995 and June 1996 were used to study the broad scale distribution patterns of anchovy eggs and larvae over the northern Aegean Sea continental shelf and the regional/inter-annual variability in growth and mortality rates of larvae. Two major spawning grounds were identified. One in the east, located in the area influenced by the Samothraki gyre (SG), in which a large amount of enriched, modified Black Sea water (BSW) is entrapped and one in the west, associated with zooplankton-rich waters in the semi-enclosed Thermaikos gulf close to several river mouths. In the NE Aegean, anticyclonic gyres generated over the continental shelf and fed by the circulating stream of BSW (like the SG) may act as retention areas for larval anchovy. In the west, the high enclosure of the Thermaikos Gulf contributes to reducing offshore dispersal. Major changes were observed in egg and larval abundance as well as larval mortality between June 1995 and June 1996 in both the western and eastern part of the continental shelf. Mean abundance of eggs and early larvae was >5 times higher in 1996 than in 1995, when waters were significantly cooler, fresher and richer in mesozooplankton. Larval survival decreased from 79 to 69% day⁻¹ in the east and from 89 to 74% day⁻¹ in the west between 1995 and 1996. Hence increased egg production was coupled with higher larval mortality during June 1996. Furthermore, a highly significant positive relationship between larval mortality (Z) and mean egg abundance (A) emerged (Z = −0154 + 0.205 log[A], r ² = 0.96, n = 7) when data from this study and a similar study in the NW Mediterranean were regressed. Mean growth rate of anchovy larvae in the study area (∼0.5 mm day⁻¹) did not differ significantly between areas/years. A marked ontogenetic change was observed in the otolith size/recent otolith growth-on-fish size relationships, which exhibited significant inflection points at ∼6 mm formalin preserved length. This change seems to coincide with performance (e.g., catchability) and behavioral changes (e.g., onset of vertical migrations) in European anchovy associated with the development of the caudal fin (the flexion stage).     

Vertical Distribution and Inventories of 239,240Pu in Deep Sea Sediments of the Mediterranean Sea (Algerian Basin, Ionian Sea)

Deep-sea sediments collected in two areas of the Mediterranean Sea having different sedimentological characteristics have been analyzed to determine present concentrations and inventories of ^239,240Pu and to study the main mechanisms controlling them. Plutonium distribution in the deepest part of a submarine canyon (Taranto Valley, Ionian Sea) is compared to that obtained in an abyssal plain (north Algerian Plain). in the latter case, sedimentation is mainly due to the sinking of biogenic particles, while in the former lateral transport of terrigenous material along the slope of the canyon can significantly contribute to sediment accumulation on the bottom.

^239,240Pu surface concentration in the canyon ranged from 0.2 to 1 Bq kg^-1 (dry weight) and this was lower in the abyssal plain. in this area, plutonium was detectable only in the first 4 cm, while in the canyon it was present down to 11-15 cm. ^239,240Pu inventories are 3 Bq m^-2 in the plain and 45-60 Bq m^-2 in the canyon, indicating considerable input of terrigenous material towards the final part of the Taranto Valley.     

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.     

Reproduction and distribution of the invasive European fanworm Sabella spallanzanii (Polychaeta: Sabellidae) in Port Phillip Bay, Victoria, Australia

 The European fanworm Sabella spallanzanii (Gmelin, 1791) was recently introduced to Port Phillip Bay and is now a conspicuous component of most benthic communities. Reproduction of the worm was investigated in a population at Queenscliff over a 2 yr period (October 1995 to October 1997) using gonadal histology. The worms are dioecious (sex ratio 1:1, n=250), and attained sexual maturity at ∼50 mm body length. Reproductive periodicity followed a distinct annual cycle, and spawning proceeded through an extended autumn/winter period. Spawning was broadly synchronous between sexes, and coincided with falling seawater temperatures and shorter day-lengths. The females were highly fecund, and >50 000 eggs were probably shed from large females (>300 mm body length) during the annual spawning period. Breeding cycles of S. spallanzanii in Port Phillip Bay are ∼6 mo out of phase with endemic populations located at similar latitudes in the northern hemisphere. The spread of S. spallanzanii within Port Phillip Bay has been monitored by divers on an annual basis since 1994. The most recent dive survey (1998) indicates that S. spallanzanii has extended its range through out the entire 2000 km² embayment, and has invaded most subtidal habitats. Quantitative estimates of S. spallanzanii abundances were highest on pier pylons (12.5 individuals m⁻², 0.5 to 7 m depths). On sediments, estimates were highest at shallow sites (0.3 m⁻², 7 m depth), but numbers declined significantly with depth (0.1 m⁻², 17 to 22 m depth). Mean worm lengths and biomass were, by contrast, significantly higher at intermediate depths (12 to 17 m) than in shallower (7 m) or deeper (22 m) locations. S. spallanzanii demonstrates a clear preference for growth in sheltered, nutrient-enriched waters, so it may not spread from Port Phillip Bay into the adjacent oceanic waters of Bass Strait; however, in view of S. spallanzanii's current high abundance, fecundity and extended spawning periodicity, there is a high risk of future range expansions, mediated by shipping, into other temperate-water ports. Received: 17 November 1998 / Accepted: 6 January 2000     

Fine-scale distribution of juvenile Atlantic and Mediterranean loggerhead turtles (Caretta caretta) in the Mediterranean Sea

Loggerhead turtles nesting in the Mediterranean Sea exhibit remarkable genetic structuring. This paper tests the hypothesis that young loggerhead turtles from different rookeries do not distribute homogeneously among the major Mediterranean foraging grounds, due to a complex pattern of surface currents. We extracted long fragments of mitochondrial DNA from 275 stranded or bycaught juvenile turtles from six foraging grounds (Catalano-Balearic Sea, Algerian basin, Tyrrhenian Sea, Adriatic Sea, northern Ionian Sea and southern Levantine Sea). We used a Bayesian mixed-stock analysis to estimate the contributions from rookeries in the Mediterranean, the North-west Atlantic and Cape Verde to the studied foraging grounds. Differences were found in the relative contribution of juvenile turtles of Atlantic and Mediterranean origin to each foraging ground. A decreasing proportion of Atlantic juveniles was detected along the main surface current entering the Mediterranean, with a high prevalence of turtles from eastern Florida in the Algerian basin and lower numbers elsewhere. In regard to the turtles of Mediterranean origin, juveniles from Libya prevailed in central and western Mediterranean foraging grounds other than the Algerian basin. Conversely, the Adriatic Sea was characterised by a large presence of individuals from western Greece, while the southern Levantine Sea was inhabited by a heterogeneous mix of turtles from the eastern Mediterranean rookeries (Turkey, Lebanon and Israel). Overall, the distribution of juveniles may be related to surface circulation patterns in the Mediterranean and suggests that fisheries might have differential effects on each population depending on the overlap degree between foraging and fishing grounds.     

The effect of mirrors on African clawed frog (Xenopus laevis) larval growth, development, and behavior

We examined the behavioral and developmental responses of Xenopus laevis larvae to their mirror images in three experiments. The mirrors allowed us to visually simulate increased density, without the tadpoles’ behavior being confounded by chemical cues from additional tadpoles. In the first experiment, we demonstrated that Xenopus tadpoles have a right eye preference for mirrors, contrary to the left eye preference of all other anuran species studied to date. This lateralized eye use disappeared, however, as tadpoles approached metamorphosis. Next, we examined how mirrored aquaria walls affected tadpole growth and development. We found that tadpoles raised in aquaria with partially mirrored walls showed depressed growth compared to tadpoles raised without mirrors, despite the fact that Xenopus larvae normally thrive when raised in visual contact with conspecifics. The tadpoles raised with mirrors had, though not significantly, proportionally larger bodies relative to their tail length (d = 0.51). This suggests that a phenotypically plastic response in body proportions was induced in these tadpoles solely by the sight of other tadpoles. The third experiment established that X. laevis tadpoles are more active in front of a mirror; i.e., they turn more often and spend more time in front of mirrored surfaces. We consider this increased activity to be an aberrant behavior of the tadpoles, which were attempting to school with their own images. We suggest that this extra activity reduced the amount of energy available for growth, accounting for the depressed growth seen in our second experiment.     

Salinity favorable for early development and gamete compatibility in two sympatric estuarine species of the genus Hediste (Polychaeta: Nereididae) in the Ariake Sea, Japan

Early development was examined, under various salinities, for two sympatric nereidid polychaetes, Hediste japonica and H. diadroma, which participated in a simultaneous reproductive swarming in an estuary of the Omuta-gawa River in Ariake Sea, Japan. The eggs of both species were isotonic to the medium of 27.5–30 psu salinity. The egg diameter in the isotonic salinity was 180–205 μm in H. japonica, and 130–160 μm in H. diadroma. Successful development of most embryos was observed in a salinity range of 22.5–30 psu in both species, while successful fertilization occurred in wider ranges of salinity, i.e., 10–34 psu in H. japonica and 10 to 30 psu in H. diadroma. In both species, free-swimming larval life started from the indistinct hatching of trochophores out of a jelly layer capsule. The lecithotrophic development appeared to run to the 4-setiger nectochaetes in H. japonica, while to 3-setiger nectochaetes in H. diadroma, resulting in a shorter pelagic larval life in H. japonica. In a comparison of larval morphology among Hediste species, we found a definite negative correlation between the prostomium width, which represents the larval size and depends on egg size, and relative length of chaetae to the prostomium width: the relative length of chaetae was the longest in H. diadroma (with the smallest egg size and long pelagic life), intermediate in H. japonica (intermediate egg size, short pelagic life), and the shortest in H. atoka (largest egg size, no true pelagic life). We also examined the possibility of hybridization between H. japonica and H. diadroma through cross-insemination experiments. The gametes of the two species were reciprocally compatible, and viable hybrid offspring were produced by the laboratory crosses. The hybrid larvae expressed intermediate phenotypes, but with a greater maternal influence in characteristics such as the relative length of chaetae and the lecithotrophic larval duration.     

Reproductive cycle,larval development,juvenile growth and population dynamics of Patiriella pseudoexigua (Echinodermata: Asteroidea) in Taiwan

A field study was conducted at Wanlitung, southern Taiwan, in 1986–1089, to determine the reproductive cycle, development mode, growth rate and population dynamics of the small seastar Patiriella pseudoexigua (Dartnall), which occurs in highly stressful and disturbed intertidal pools in this area. An inverse relationship between gonad index and pyloric-caccum index was only recorded immediately prior to spawning. A short, well-synchronized seasonal spawning occurs in October. When reared at 25 °C, lecithotrophic larvae develop directly, lack a bipinnaria stage, and metamorphose completely on the seventh day after fertilization. The growth curves of field juveniles are linear, those of laboratory-reared juveniles are sigmoid. Juveniles appear in tide pools in spring-early summer of each year. Adults spawn mainly in late fall, enabling spawning to occur in time for the larvae to benefit from the environmentally favorable winter season. Populations in high-tidal pools decrease in later summer, but remain more stable in lowtidal pools and lagoons.     

Comparative biometry of Eunicella singularis (Gorgonian) sclerites at East Mediterranean Sea (North Aegean Sea, Greece)

The two forms of Eunicella singularis sclerites, spindles and clubs, were examined in two sites in East Mediterranean Sea, Greece. The comparison of the sclerites size was based on the hydrodynamic profile of the sites (the site of Arethoussa was exposed and the Phidonissi site was sheltered), on the sites bathymetry (AR 5–8 m, AR 9–13 m, PH 5–10 m, PH 11–13 m) and on the position occupied by the sclerites on the gorgonian colonies (top, middle and base). With the usage of image system analysis the following microfeatures were measured or calculated: projection, length, width, elongation factor, compactness factor, length × width and length/width. The present study confirms the hypothesis that spindles are involved in the recovering of the gorgonian colonies and clubs contribute to the flexion capacity of the colonies. The spindles of the shallow E. singularis colonies (AR 5–8 m) were significantly bigger (length, width) compared with the others, in order to assure the best recovery of the colonies after their flexion, induced by higher water current velocity in shallow water. The clubs occupy the intern part of the mesoglea and their size (length, width) increased in colonies from the shallow exposed site in order to facilitate the gorgonian flexion. The present study demonstrates the gorgonian adaptation (in a microscopic scale) to the environmental pressure of hydrodynamic conditions.     

Genetic consequences of long larval life in the starfish Linckia laevigata (Echinodermata: Asteroidea) on the Great Barrier Reef

Gene flow between populations of the asteroid Linckia laevigata (Linnaeus) was investigated by examining over 1000 individuals collected from ten reefs throughout the Great Barrier Reef (GBR), Australia, for genetic variation at seven polymorphic enzyme loci. Despite geographic separations in excess of 1000 km, Nei's unbiased genetic distance (0 to 0.003) and standardised genetic variation between populations (F _ST) values (mean 0.0011) were small and not significant. Genetic homogeneity among L. laevigata populations is consistent with the long-distance dispersal capability of its 28 d planktonic larval phase, and is greater than that observed for other asteroid species, including another high-dispersal species, Acanthaster planci, which has a 14 d larval phase. Variation within populations was also higher than previously recorded for asteroids (mean heterozygosity=0.384; number of alleles per locus ranged from 5.1 to 6.0 in each population). Among asteroids, dispersal ability is positively correlated with gene flow and levels of variation, and negatively correlated with levels of differentiation.