Differential ingestion of zooplankton by four species of bivalves (Mollusca) in the Mali Ston Bay,Croatia

This study provides information about differences in composition of ingested zooplankton amongst bivalve species coexisting in the same area in a period from May 2009 to December 2010. The study was conducted at the Mali Ston Bay (42°51′ N, 17°40′ E)—the most important bivalve aquaculture area in the eastern Adriatic Sea. Stomach content analysis was performed on cultured species—Ostrea edulis and Mytilus galloprovincialis, and commercially important bivalve species from their natural environment—Modiolus barbatus and Arca noae. Results confirmed carnivory in bivalves, both from natural and cultured populations, but cultured species had higher numbers of zooplankters than those living on the seabed. The most abundant taxa were bivalve larvae, followed by tintinnids, copepods, unidentified eggs and gastropod larvae. Recorded numbers of bivalve larvae in M. galloprovincialis stomach were the highest so far reported and show that mussels impact the availability of natural spat.     

<Emphasis Type="Italic">Cochlodinium polykrikoides</Emphasis> blooms and clonal isolates from the northwest Atlantic coast cause rapid mortality in larvae of multiple bivalve species

Globally, many commercial bivalve populations have declined in recent decades. In addition to overharvesting and habitat loss, the increasing frequency and intensity of harmful algal blooms (HABs) are likely to contribute to bivalve losses, particularly in cases where blooms negatively impact larval stages. This paper reports on the lethal effects of clonal cultures and blooms of Cochlodinium polykrikoides from the US Atlantic coast on the larvae of three species of commercially and ecologically valuable bivalves: the Eastern oyster (Crassostrea virginica), the bay scallop (Argopecten irradians), and the Northern quahog (hard clam; Mercenaria mercenaria). Both cultures and blooms of C. polykrikoides were highly toxic to all three species of bivalve larvae causing 80–100% mortality during 24- to 72-h exposures at concentrations of 1–2 × 10³ cells ml⁻¹. Toxicity was dependent on cell densities, growth stage of C. polykrikoides (i.e. cultures in exponential stage growth were more toxic than later stages), exposure time of larvae to cells (i.e. longer exposure caused higher mortality), the age of larvae (i.e. younger larvae were more sensitive), and the relative abundance of C. polykrikoides (i.e. the presence of other microalgae decreased toxicity). Free radical-scavenging enzymes (peroxidase and catalase) and the removal of C. polykrikoides cells (i.e. culture filtrate) significantly increased larval survival suggesting toxicity is maximized by contact with live cells and may involve labile toxins bound by these compounds including e.g. reactive oxygen species. The toxicity of C. polykrikoides to bivalve larvae was generally more severe than other HAB species (e.g. Karenia brevis, Karlodinium veneficum, Alexandrium tamarense, Prorocentrum minimum). Since the bivalves in this study spawn in the months when C. polykrikoides blooms on the east coast of North America, these results suggest that these blooms may have detrimental effects on efforts to restore these already diminished populations.     

Settlement of <Emphasis Type="Italic">Macoma balthica</Emphasis> larvae in response to benthic diatom films

The role of multi-species benthic diatom films (BDF) in the settlement of late pediveliger larvae of the bivalve Macoma balthica was investigated in still-water bioassays and multiple choice flume experiments. Axenic diatom cultures that were isolated from a tidal mudflat inhabited by M. balthica were selected to develop BDF sediment treatments characterized by a different community structure, biomass, and amount of extracellular polymeric substances (EPS). Control sediments had no added diatoms. Although all larvae settled and initiated burrowing within the first minute after their addition in still water, regardless of treatment, only 48–52% had completely penetrated the high diatom biomass treatments after 5 min, while on average 80 and 69% of the larvae had settled and burrowed into the control sediments and BDF with a low diatom biomass (<3.5 μg Chl a g⁻¹ dry sediment), respectively. The percentage of larvae settling and burrowing into the sediment was negatively correlated with the concentration of Chl a and EPS of the BDF. This suggests higher physical resistance to bivalve penetration by the BDF with higher diatom biomass and more associated sugar and protein compounds. The larval settlement rate in annular flume experiments at flow velocities of 5 and 15 cm s⁻¹ was distinctly lower compared to the still-water assays. Only 4.6–5.8% of the larvae were recovered from BDF and control sediments after 3 h. Nonetheless, a clear settlement preference was observed for BDF in the flume experiments; i.e., larvae settled significantly more in BDF compared to control sediments irrespective of flow speed. Comparison with the settlement of polystyrene mimics and freeze-killed larvae led to the conclusion that active selection, active secondary dispersal and, at low flow velocities (5 cm s⁻¹), passive adhesion to the sediment are important mechanisms determining the settlement of M. balthica larvae in estuarine biofilms.     

Lethal effects of Northwest Atlantic Ocean isolates of the dinoflagellate, Scrippsiella trochoidea, on Eastern oyster (Crassostrea virginica) and Northern quahog (Mercenaria mercenaria) larvae

The thecate dinoflagellate Scrippsiella trochoidea is a cosmopolitan, bloom-forming alga that has been generally considered non-toxic. Here, we report that environmentally relevant cell densities (10⁴ cells mL⁻¹) of Scrippsiella trochoidea strains isolated from the Northwest Atlantic Ocean caused 100% mortality in Eastern oyster (Crassostrea virginica) larvae during 3-day exposures while parallel control larvae exhibited 100% survival. S. trochoidea also exhibited lethal effects on Northern quahog (Mercenaria mercenaria) larvae (70% mortality during 3-day exposure) but were non-toxic to juvenile fish (Cyprinodon variegates). The cultures of S. trochoidea were more lethal to Northern quahog larvae than ten other species of harmful algae, including the highly toxic species Cochlodinium polykrikoides. Scrippsiella trochoidea cultures within later stages of growth were more toxic than exponential growth stages to bivalve larvae, and the toxicity was dose dependent. Furthermore, toxicity was maintained in the cultures that were sonicated, boiled, and frozen as well as in resuspended residues of the culture but was significantly lower in cell-free culture media. Collectively, these results suggest that S. trochoidea causes mortality in bivalve larvae through a physicochemical rather than strictly chemical mechanism, such as clogging of larval feeding apparatuses by materials produced by S. trochoidea (e.g., lipids, extracellular polysaccharides, and/or cell debris) which accumulate as cells in culture or blooms age. This is the first report of the lethal effects of Scrippsiella trochoidea on shellfish larvae.     

Single-step nested multiplex PCR to differentiate between various bivalve larvae

A nested multiplex polymerase chain reaction (PCR) assay has been developed that allows the discrimination between six bivalve larvae common to Danish coastal waters (Cerastoderma edule, Macoma balthica, Mytilus edulis, Spisula subtruncata, Ensis americanus and members of the order Myoida). This assay involves the simultaneous use of a pair of general universally targeted 18S rRNA gene primers, five specific 18S rRNA gene targeted oligonucleotide primers internal (nested) to the universal primer pair and one species-specific primer that is not nested (Mya). The specificity of each primer was evaluated in silico, empirically, and verified further by sequencing of amplification products from single larvae collected from plankton samples. Identification of individually isolated bivalve larvae from plankton samples was based on the size of the PCR product produced by the specific primers after visualisation by agarose gel electrophoresis. Preliminary studies indicated that this method was suitable for use with freshly collected and preserved larvae, and is therefore suitable for field application.Communicated by M. Kühl, Helsingør     

Life history characteristics of a coral reef sponge

Study of the life history characteristics of the common Red Sea sponge Mycale fistulifera (Poecilosclerida: Demospongiae) reveals several traits which may categorize it as an opportunistic species: (1) at least part of the population is reproductively active throughout the year, providing a continuous supply of larvae for settlement; (2) sponges may produce and release a large number (152±39 larvae d^-1) of brooded larvae; (3) released larvae are capable of fast settlement and metamorphosis (minutes to 30 hours post-release); (4) the turnover of the population is high and population size varies with time. From 48 initial sponges plus 94 recruits, only 25 remained alive after 14 months of observation; (5) most members of the population (>70%) have a small body size (<30 cm²); (6) sexual maturity may occur at an early age. These traits facilitate continuous establishment of M. fistulifera in new spaces on the reef; (7) M. fistulifera, preferred substrate is another opportunistic species, the coral Stylophora pistillata which it overgrows.     

Spawning and development of three coral-associated Lithophaga species in the Red Sea

Lithophaga date mussels from three species (L. lessepsiana, L. simplex and L. purpurea) were removed from their stony coral hosts in the Red sea at Eilat, Israel. Spawning, observed in the laboratory on several occasions during 1987–1988, appeared to be closely tied to lunar periods, occurring primarily during the last quarter and the new moon. Embryonic and larval development was typical of that described for other mytilids and, except for pigmentation differences, which could be discerned during embryogenesis, the developmental stages of the three species were indistinguishable. Development to the pediveliger stage took 3 to 4 wk in standard culture conditions, but raising the temperature to 27.5 °C increased the growth rate of larvae of L. lessepsiana by as much as three-fold, so that the pediveliger stage was attained in 16 d. Larvae resulting from spawning by L. simplex adults removed from the coral Astreopora myriophthalma grew significantly faster in culture than larvac from adults removed from the coral Goniastrea pectinata (comparison of slopes, p<0.05). The latter individuals showed a 6-d growth plateau at the early umbone stage. Metamorphically competent larvae were capable of delaying metamorphosis for up to 4mo, which would allow an extended period for dispersion and would increase the chance of finding a suitable substratum in the natural environment.     

Effects of Lunatia heros predation on the population dynamics of Mya arenaria and Macoma balthica in Maine,USA

The interaction between the naticid snail predator Lunatia heros and 2 iteroparous, infaunal, intertidal bivalves was investigated in Lubec, Maine, USA. The Mya arenaria population consists primarily of young, small individuals. M. arenaria survivorship is low when young (3.5% yr^-1 for the first 5 yr), then increases. M. arenaria can attain a length of 60 mm, but it is susceptible to L. heros attack only until it is 30 mm long. It delays reproduction until it is 20 mm long (3.8 yr) and diverts its resources instead into rapid early growth (4.9 mm yr^-1 for the first 5 yr). The Macoma balthica population has a larger proportion of older individuals than does that of M. arenaria. Survivorship is higher for M. balthica than for M. arenaria (76.3% yr^-1 for the first 5 yr). Unlike M. arenaria, M. balthica attains a final length of only 25 mm and all sizes are susceptible to L. heros attack. M. balthica grows slowly (2.7 mm yr^-1 for the first 5 yr) and diverts its resources into earlier reproduction at a length of 10 mm (2.9 yr). These contrasting life-history patterns and the possible relationship between bivalve resource allocation and refuges from predation are discussed.     

Factors influencing the abundance of the seven-spined morphotype of Spirobranchus polycerus (Schmarda), (Serpulidae), on upright blades of the hydrozoan coral,Millepora complanata

This study, on the seven-spined morphotype of Spirobranchus polycerus, a serpulid polychaete commensal with the hydrozoan coral Millepora complanata Lamarck examines live worm abundance, net recruitment and the probability of mortality, on single blades of coral at four Barbados fringing reefs, Heron Bay (HB), Greensleeves (G), Sandridge (S) and Six Men's Bay (SMB). Variation in the number of worms blade^-1 over the part 5 to 12 yr, is explained largely by variation in mortality at HB, and largely by variation in recruitment at SMB. At G and S the relative importance of these two factors appears to have shifted with time. A smaller number of worms blade^-1 at SMB than at HB, G or S, may be a consequence of a recruitment limited to the past 1 to 4 yr, which, in turn, may partly explain the interdependence of effects of reef and blade size class on recruitment and mortality. The relationship between level of recruitment and blade base perimeter suggests that the availability of recruits has been consistently high at HB over the past 5 to 12 yr and has increased at G and S in the past 1 to 4 yr. The situation is consistent with a worm population gradually extending northward, dependent on a pool of larvae to the south and limited by a tendency to lose larvae in the north west drift. Because of periodic destruction of tall coral blades in heavy seas, mortality in the seven-spined S. polycerus is, in part, a result of the commensal relationship between the polychaete and the coral. Selection for simultaneous hermaphroditism may be a response to the short life-span of the seven-spined form of S. polycerus and the isolation of breeding individuals resulting from the natural destruction and discontinuous distribution of the host coral.     

Energy relations of the bivalve Mercenaria mercenaria on an intertidal mudflat

Measurements of the filtration rate and oxygen consumption of the bivalve Mercenaria mercenaria (L.) have been made using flowing water systems, and then combined with growth and survivorship data to construct an energy budget for a tidal-flat population in Southampton Water. Estimated consumption (1292 kcal m^-2 year^-1) is mainly deposited as faeces and pseudofaeces (759 kcal) or excreted (160 kcal); 29% (374 kcal) is assimilated, of which 241 kcal are used for respiration, 72 kcal for flesh production and 61 kcal for gamete production. M. mercenaria contributes significantly to other trophic groups in Southampton Water; predators remove an estimated 55 kcal m^-2 year^-1, and 829 kcal pass to the scavenger/decomposer chain.     

Toxicological and larvicidal activities of Alzanzalakhet,Melia azedarach against cucurbit fly,Dacus ciliatus at Hail Province in Saudi Arabia

Alzanzalakhet, Melia azedarach L. (Sapindales: Meliaceae) is a common medicinal plant which is found in Hail desert in Saudi Arabia and traditionally used for various purposes. In this study, toxin effectiveness of the ethanolic, acetonic, and aqueous extracts of M. azedarach leaves was tested to determine a larvicidal activity against the cucurbit fly, Dacus ciliatus (Loew) (Diptera: Tephritidae). Different treatments with M. azedarach extracts varied significantly in their impact on mortality of D. ciliatus larvae. Results showed that all tested extracts in the three solvents exerted a positive pattern for reducing larvae of D. ciliatus. However, the ethanolic M. azedarach extract was the most potent compared with other tested solvents. The highest mortality of the D. ciliatus larvae was shown to be 94%, 74%, and 68% when the concentration was 1?×?10⁵?ppm for ethanolic, acetonic, and aqueous extracts, respectively. In contrast, the concentration 1?×?10¹?ppm resulted in the lowest mortality of D. ciliatus larvae at 40%, 34%, and 12% for ethanolic, acetonic, and aqueous extracts, respectively. Overall, the results of this study suggest that the ethanolic extract of M. azedarach leaves may be used as a sustainable method for controlling D. ciliatus larvae.     

Slowest of the slow: latitudinal insensitivity of burrowing capacity in the bivalve <Emphasis Type="Italic">Laternula</Emphasis>

Low temperature limits the rate of biochemical reactions and aerobic scopes of cold water ectotherms. To compensate for this limiting effect, animals living in cold environments often possess physiological or morphological adaptations to maintain vital functions. Cross-latitudinal comparison of aerobic capacities is one method to test which factors constrain activity in thermally distinct environments particularly when congeneric studies are carried out on related species with conservative ecology and habitat. Burrowing is a major aerobic activity of bivalve molluscs that is described here for the first time for the tropical mangrove species Laternula truncata and Laternula boschasina and then compared with their Antarctic congener Laternula elliptica. About 80% of L. truncata (16.3–46.1 mm shell length) and 63% of L. boschasina (11.3–27.7 mm shell length) buried within 24 h at 28°C. The burrowing rate index (BRI = [³√wet weight/time to bury]×10⁴) ranged between 1.1 and 20.2 for L. boschasina and 1.1–32.9 for L. truncata. These values are 2–3 orders of magnitude less than other tropical bivalve molluscs and are amongst the lowest recorded for any bivalve. Comparisons with the Antarctic L. elliptica showed little or no differences in BRI (Q ₁₀ of 1.0–1.2 for specimens of the same size). This is contrary to the general pattern over a wide range of bivalves, where BRI increases with a Q ₁₀ of between 2.9 and 6.4 between high latitudes and the equator. L. elliptica has 25–30% longer relative foot length than tropical congeners of the same size, which could be a morphological adaptation compensating for reduced burrowing speeds in a colder environment. Burrowing rates within the genus Laternula could, however, also be maintained by differing habitat, ecological and physiological constraints on burrowing capability.     

Assemblage shift following population collapse of a non-indigenous bivalve in an urban lagoon

The quahog Mercenaria mercenaria has been introduced repeatedly to the Pacific coast of North America, but only one population is known to have become established. In the 1970s, the population of M. mercenaria at Colorado Lagoon, in Los Angeles County, California (33^o46′16″N, 118^o08′05″W), was estimated at more than 300,000 individuals. To determine the current status of this non-indigenous species (NIS), in 2009, we sampled 57 intertidal and 20 shallow subtidal plots, identifying and quantifying collected bivalves. No quahogs were found among the 2,490 living bivalves in our plots, though two were found intertidally outside of our plots. The M. mercenaria population has thus collapsed since 1980, but the native community has not recovered. Six of the fourteen living bivalve species we encountered were NIS; three are new records for the location, including the clam Venerupis philippinarum, which made up 87.6% of collected individuals. Though M. mercenaria is likely on its way to extinction on the US Pacific coast, the bivalve assemblage at this location remains heavily dominated by NIS.     

The relative contribution of dinoflagellate photosynthesis and stored lipids to the survivorship of symbiotic larvae of the reef-building corals

The long-distance dispersal of larvae provides important linkages between populations of reef-building corals and is a critical part of coral biology. Some coral planulae have symbiotic dinoflagellates (Symbiodinium spp.) that probably provide energy in addition to the lipids provisioned within the egg. However, our understanding of the influence of symbionts on the energy metabolism and survivorship of planulae remains limited. This study examines the relative roles of symbiotic dinoflagellate photosynthesis and stored lipid content in the survivorship of the developing stages of the corals Pocillopora damicornis and Montipora digitata. We found that survivorship decreased under dark conditions (i.e. no photosynthetic activity) for P. damicornis and M. digitata at 31 and 22 days after release/spawning, respectively. The lipid content of P. damicornis and M. digitata planulae showed a significant decrease, at a higher rate, under dark conditions, when compared with light conditions. When converted to energy equivalents, the available energy provided by the depletion of lipids could account for 41.9 and 84.7% of larval metabolism for P. damicornis (by day 31) and 38.4 and 90.1% for M. digitata (by day 21) under light and dark conditions, respectively. This finding indicates that not all energy requirements of the larvae are met by lipids: energy is also sourced from the photosynthetic activities of the symbiotic dinoflagellates within these larvae, especially under light conditions. In addition, the amounts of three main lipid classes (wax esters, triglycerides, and phospholipids) decreased throughout the experiment in the planulae of both species, with the wax ester content decreasing more rapidly under dark conditions than under light conditions. The observations that the planulae of both species derive considerable amounts of energy from wax esters, and that symbiotic dinoflagellates enable larvae to use their stores at lower rates, suggested that symbiotic dinoflagellates have the potential to extend larval life under light conditions.     

Hybridization within the species complex of the scleractinan coral Montastraea annularis

The morphologically variable reef coral previously known as Montastraea annularis (Ellis and Solander, 1786) has recently been separated into three species based on differences in morphology, behavior, allele frequencies and some life-history traits of Panamanian specimens. To further investigate the proposed reclassification and its conformity to the biological species concept we conducted reciprocal intra- and inter- specific fertilization experiments with gametes from each of the three species on Florida reefs. With one exception, self-fertilization rates were very low or zero. Within-species crosses resulted in production of planulae, as did all inter-species (hybrid) crosses, but there was much variation in fertilization success within each type of cross. In an experiment with separated gametes, hybrid crosses between M. annularis and M. franksi produced more larvae than within-species crosses for each species. Hybridization crosses between M. faveolata and the other two species produced fewer larvae than did within- M. faveolata crosses in the experiment with separated gametes, but many larvae resulted when the hybridizations were performed by mixing entire gamete bundles. Additional observations showed that M. franksi had 20% larger eggs and fewer eggs per gamete bundle than did the other two species and that it consistently spawned 1 to 1.5 h before the others, a potential temporal barrier to hybridization. These results indicate that there is no inherent pre-zygotic barrier to cross-fertilization among the three morphological species, although post-zygotic survival and fertility remain to be determined. The adherence of the proposed reclassification to the biological species concept requires further examination. Received: 16 April 1997 / Accepted: 17 June 1997     

Ultraviolet radiation effects on the behavior and recruitment of larvae from the reef coral Porites astreoides

We tested the rarely considered hypothesis that the ultraviolet portion (UVR, 280–400 nm) of the light spectrum affects patterns of recruitment in reef-building corals. The premise for this hypothesis rests in the fact that biologically relevant intensities of UVR penetrate to considerable depths (>24 m) in the clear waters surrounding many coral reefs, and that reef organisms allocate substantial resources to prevent and repair UVR damage. The ability of larvae spawned by the brown morph of the Caribbean coral, Porites astreoides, to detect and avoid UVR was assessed in petri dishes where one-half of the dish was shielded from UVR and the other exposed. Observations made every 30 min between 10:30 and 13:30 h showed significantly higher densities of larvae swimming in regions shielded from UVR. To determine how this behavior affects settlement patterns, larvae collected from P. astreoides adults at 18 m depth were released into chambers deployed at 17 m depth where they were given a choice of three different light regions in which to settle: PAR (PAR=400–700 nm), PAR+UVAR (UVAR=320–400 nm), and PAR+UVAR+UVBR (UVBR=280–320 nm). At the end of the experiment, greater numbers of P. astreoides larvae had settled in the region of the tube where UVR was reduced than would be expected if dispersion were random. To our knowledge, this is the first demonstration in any reef-building coral species that planula larvae can detect UVR and that it affects their choice of a settlement site. These results indicate that the capacity to detect and avoid habitats with biologically damaging levels of UVR may be one factor contributing to the successful recruitment of coral larvae.     

Ingestion of bivalve larvae by<Emphasis Type="Italic"> Mytilus edulis</Emphasis>: experimental and field demonstrations of larviphagy in farmed blue mussels

Ingestion of bivalve larvae by Mytilus edulis was investigated. Laboratory experiments revealed that ~ 90% of bivalve larvae offered to mussels was ingested and apparently fully digested. The shell of the bivalve larvae offered no protection against digestive processes, resulting in high larval mortality once inside the stomach. Stomach content analysis (September 2001–January 2003) showed that bivalve larvae were ingested by farmed mussels year-round, with the exception of March 2002. Numbers of ingested larvae were highest in October 2001 and May 2002, which coincides with known spawning times of farmed mussels in Ireland. Mussels ingested a large size-range of bivalve larvae, suggesting that all stages of the bivalve life cycle are vulnerable to predation. It is suggested that adult bivalves routinely filter larvae from the surrounding water and that, given the high biomass of mussels present in mussel farms, filtration by adult bivalves significantly reduces numbers of bivalve larvae in nearby waters.Communicated by J.P. Thorpe, Port Erin     

Rafting of reef corals and other organisms at Kwajalein Atoll

Studies conducted at Kwajalein Atoll (9°N; 168°E) in early 1988 reveal that marine organisms are commonly rafted into the area on drift pumice, drift wood and other flotsam. Coralskeletons on pumice provide the most useful quantitative data because they persist after rafted motile organisms have departed and rafted sessile forms have decomposed or been scavenged. The estimated minimum number of pumice fragments carrying corals into Kwajalein Atoll during its geological history is on the order of 10⁹, with a more realistic estimate in excess of 10¹¹. The estimated number of coral colonies rafted into the atoll would probably have to be increased several-fold if rafting on floating organic materials such as wood, charcoal, nuts and seeds could be determined. In the present study, a typical sample of beached pumice from Kwajalein Atoll contained 10³ coral colonies per m³ of bulk pumice fragments. Major pumice rafting episodes frequently result from volcanic eruptions. Estimates based on the observed K wajalein coral fouling rate and literature values for drift pumice production suggest that 10⁵ colonies could be rafted through the tropics by the smallest reported pumice-producing events and up to 10¹² colonies for large events. In the present study, most of the rafted corals that were recovered belong to species of the genus Pocillopora, but speccics of Porites and Millepora were also collected. Analysis of surface-current data and reported drift patterns for pumice, logs, drift bottles and wrecks suggests that movement of rafted corals and drifting larvae is predominantly from peripheral areas of low coral-species diversity into centers of high coral diversity. Coral diversity centers might be viewed as areas of coral species accumulation rather than centers of coral species origin.     

Accumulation of polychlorinated biphenyls in the cockle Cerastoderma edule and the tellin Macoma balthica

Two species of marine bivalve, Cerastoderma edule (Linnaeus) and Macoma balthica (Linnaeus), were exposed to particles contaminated with three commercial polychlorinated biphenyl (PCB) mixtures, Aroclors^® 1242, 1254 and 1260. The accumulation of PCB in bivalve tissue was monitored for periods of up to 40 days. In addition to measuring total PCB concentrations, which reached 60 ppm of Aroclor 1242 in M. balthica, the fate of individual PCB homologues of different chlorine content, ranging from 2 to 8 chlorine atoms per biphenyl molecule, was also determined. Selective accumulation of PCBs with 5 chlorine atoms per biphenyl molecule occurred, whilst isomers of very low and very high molecular weights were accumulated at much slower rates.     

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.