Morphological process of vertical transmission of photosymbionts in the colonial ascidian <Emphasis Type="Italic">Trididemnum miniatum</Emphasis> Kott, 1977

In obligate symbioses between didemnid ascidians and prokaryotic algae (cyanobacteria), the larvae of the host ascidian usually possess Prochloron cells that are acquired from their mother colony. The process of vertical transmission of the photosymbionts has been histologically described in several species harboring Prochloron in the common cloacal cavity, where Prochloron cells are attached to the hairy surface of the tunic of the larvae during or just before larval spawning. Since the process has never been described in species harboring the photosymbionts in the tunic of the colony, here we describe the histological and ultrastructural process of vertical transmission in a photosymbiotic didemnid, Trididemnum miniatum. No photosymbionts were associated with gametes of the ascidian. Prochloron cells appeared in the epithelial pouch enclosing the embryo and were then incorporated in the tunic of embryos in late embryonic stages. Although Prochloron cells in the tunic of the colony were rarely associated with the host cells, some amoeboid-shaped tunic cells containing Prochloron cells were occasionally found around the epithelial pouch. Therefore, the host cell is thought to be a vehicle transporting the photosymbionts in the tunic of the colony to the tunic of the embryos. The photosymbiont cells were directly embedded in the tunic of the larval trunk. They were not contained in the host cells, as are those in the tunic of the colony. These observations revealed that the mechanism of vertical transmission in T. miniatum is very different from that in didemnids harboring Prochloron in the common cloacal cavity of the colonies, such as Trididemnum cyclops. The mechanisms of symbiont transmission are diverse even within the genus Trididemnum.     

Temporal and spatial infection dynamics indicate recognition events in the early hours of a dinoflagellate/coral symbiosis

The obligate symbiotic relationship between dinoflagellates, Symbiodinium spp. and reef building corals is re-established each host generation. The solitary coral Fungia scutaria Lamarck 1801 harbors a single algal strain, Symbiodinium ITS2 type C1f (homologous strain) during adulthood. Previous studies have shown that distinct algal ITS2 types in clade C correlate with F. scutaria—Symbiodinium specificity during the onset of symbiosis in the larval stage. The present study examined the early specificity events in the onset of symbiosis between F. scutaria larvae and Symbiodinium spp., by looking at the temporal and spatial infection dynamics of larvae challenged with different symbiont types. The results show that specificity at the onset of symbiosis was mediated by recognition events during the initial symbiont—host physical contact before phagocytosis, and by subsequent cellular events after the symbionts were incorporated into host cells. Moreover, homologous and heterologous Symbiodinium sp. strains did not exhibit the same pattern of localization within larvae. When larvae were infected with homologous symbionts (C1f), ~70% of the total acquired algae were found in the equatorial area of the larvae, between the oral and aboral ends, 21 h after inoculation. In contrast, no spatial difference in algal localization was observed in larvae infected with heterologous symbionts. This result provides evidence of functional differences among gastrodermal cells, during development of the larvae. The cells in the larval equator function as nutritive phagocytes, and also appear to function as a region of enhanced symbiont acquisition in F. scutaria.     

Elevated temperature impairs onset of symbiosis and reduces survivorship in larvae of the Hawaiian coral, <Emphasis Type="Italic">Fungia scutaria</Emphasis>

Many corals obtain their obligate intracellular dinoflagellate symbionts from the environment as larvae or juveniles. The process of symbiont acquisition remains largely unexplored, especially under stress. This study addressed both the ability of Fungia scutaria (Lamarck 1801) larvae to establish symbiosis with Symbiodinium sp. C1f while exposed to elevated temperature and the survivorship of aposymbiotic and newly symbiotic larvae under these conditions. Larvae were exposed to 27, 29, or 31°C for 1 h prior to infection, throughout a 3-h infection period, and up to 72 h following infection. Exposure to elevated temperatures impaired the ability of coral larvae to establish symbiosis and reduced larval survivorship. At 31°C, the presence of symbionts further reduced larval survivorship. As sea surface temperatures rise, coral larvae exposed to elevated temperatures during symbiosis onset will likely be negatively impacted, which in turn could affect the establishment of future generations of corals.     

Onset of symbiosis and distribution patterns of symbiotic dinoflagellates in the larvae of scleractinian corals

The establishment of symbiosis in early developmental stages is important for reef-building corals because of the need for photosynthetically derived nutrition. Corals spawn eggs and sperm, or brood planula larvae and shed them into the water. Some coral eggs or planulae directly inherit symbiotic dinoflagellates (Symbiodinium spp.) from their parents, while others acquire them at each generation. In most species examined to date, the larvae without dinoflagellates (aposymbiotic larvae) can acquire symbionts during the larval stage, but little is known regarding the timing and detailed process of the onset of symbiosis. We examined larval uptake of symbiotic dinoflagellates in nine species of scleractinian corals, the onset of symbiosis through the early larval stages, and the distribution pattern of symbionts within the larval host, while living and with histology, of two acroporid corals under laboratory conditions. The larvae acquired symbiotic dinoflagellates during the planktonic phase in all corals examined which included Acropora digitifera, A. florida, A. intermedia, A. tenuis, Isopora palifera, Favia pallida, F. lizardensis, Pseudosiderastrea tayamai, and Ctenactis echinata. The larvae of A. digitifera and A. tenuis first acquired symbionts 6 and 5 days after fertilization, respectively. In A. digitifera larvae, this coincided with the formation of an oral pore and coelenteron. The number of symbiotic dinoflagellates increased over the experimental periods in both species. To test the hypothesis that nutrients promotes symbiotic uptake, the number of incorporated dinoflagellates was compared in the presence and absence of homogenized Artemia sp. A likelihood ratio test assuming a log-linear model indicated that Artemia sp. had a significantly positive effect on symbiont acquisition. These results suggest that the acquisition of symbiotic dinoflagellates during larval stages is in common with many coral species, and that the development of both a mouth and coelenteron play important roles in symbiont acquisition.     

N2 fixation (nitrogenase activity) attributable to a specific Prochloron (Prochlorophyta)-ascidian association in Palau,Micronesia

Light-mediated nitrogenase activity (NA) consistently occurred in one of four species of tropical marine ascidians (sea squirts) hosting the symbiotic alga Prochloron (prochlorophyta) among reef habitats located in Palau, Micronesia. NA was limited to intact colonies of Lissoclinum patella exclusively colonized by cells of Prochloron. Neither isolated viable cells of Prochloron nor ascidians free of Prochloron revealed NA, indicating a strong dependence on the intact symbiosis for creating conditions conductive to N₂ fixation. The confinement of NA to L. patella may be related to both the oligotrophic habitat and the large colony size of this protochordate species. This is the first report of NA attributable to symbiotic alga residing in either an aquatic or a terrestrial animal.     

Association of bacteria with larvae of the gutless protobranch bivalve Solemya reidi (Cryptodonta: Solemyidae)

Rod-shaped bacteria were consistently observed by transmission electron microscopy in the locomotory test of larvae and in the perivisceral cavity of post-larvae of Solemya reidi, a gutless protobranch bivalve known to possess intracellular chemoautotrophic bacterial symbionts in the adult gill. Bacteria develop within granular vesicles in the larval test, where they either remain to be ingested at metamorphosis, or are released into the space separating the test and embryo, to be subsequently ingested through the larval mouth. In either case, bacteria lie within the perivisceral cavity following metamorphosis. Bacteria were not seen either in or on gametes or in gills of juveniles. It is hypothesized that these bacteria represent a transmission stage of the gill symbionts present in adult S. reidi and are not evident in gametes or gills of juveniles due to cryptic packaging within granular vesicles. Perpetuation of this symbiosis would therefore be assured through vertical transmission, as is typical of other marine invertebrate-bacteria endosymbioses.Harbor Branch Oceanographic Institution Contribution No. 602     

Biochemical composition,energy content and chemical antifeedant and antifoulant defenses of the colonial Antarctic ascidian<Emphasis Type="Italic"> Distaplia cylindrica</Emphasis>

The colonial ascidian Distaplia cylindrica occurs both as scattered individual colonies or in gardens of colonies in fine-grained soft substrata below 20 m depths off Anvers Island along the Antarctic Peninsula. Individual colonies, shaped as tall rod-like cylinders and anchored in the sediments by a bulbous base, may measure up to 7 m in height. D. cylindrica represent a considerable source of materials and energy for prospective predators, as well as potential surface area for fouling organisms. Nonetheless, qualitative in situ observations provided no evidence of predation by sympatric predators such as abundant sea stars, nor obvious biofouling of colony surfaces. Mean energy content of whole-colony tissue of D. cylindrica was relatively high for an ascidian (14.7 kJ g^–1 dry wt), with most of this energy attributable to protein (12.7 kJ g^–1 dry wt). The sympatric omnivorous sea star Odontaster validus consistently rejected pieces of D. cylindrica colonies in laboratory feeding assays, while readily ingesting similarly sized alginate food pellets. Feeding deterrence was determined to be attributable to defensive chemistry, as colonies of D. cylindrica are nutritionally attractive and lack physical protection (conspicuous skeletal elements or a tough outer tunic), and O. validus display significant feeding-deterrent responses to alginate food pellets containing tissue-level concentrations of organic extracts. In addition, high acidity measured on outer colony surfaces (pH 1.5) as well as homogenized whole-colony tissues (pH 2.5) are indicative of surface sequestration of inorganic acids. Agar food pellets prepared at tissue levels of acidity resulted in significant feeding deterrence in sea stars. Thus, both inorganic acids and secondary metabolites contribute to chemical feeding defenses. D. cylindrica also possesses potent antifoulant secondary metabolites. Tissue-level concentrations of hydrophilic and lipophilic extracts caused significant mortality in a sympatric pennate diatom. Chemical feeding deterrents and antifoulants are likely to contribute to the abundance of D. cylindrica and, in turn, play a role in regulating energy transfer and community structure in benthic marine environments surrounding Antarctica.Communicated by P.W. Sammarco, Chauvin     

Three party symbiosis: acoelomorph worms,corals and unicellular algal symbionts in Eilat (Red Sea)

Epizoic worms were found to occur on certain coral colonies from reefs off the coast of Eilat (Red Sea). We identified 14 coral species infested by acoelomorph worms at a depth range of 2–50 m. The host corals were all zooxanthellate and included both massive and branching stony corals and a soft coral. Worms from all hosts were identified as belonging to the genus Waminoa and contained two distinct algal symbionts differing in size. The smaller one was identified as Symbiodinium sp. and the larger one is presumed to belong to the genus Amphidinium. Worm-infested colonies of the soft coral, Stereonephthya cundabiluensis, lacked a mucus layer and exhibited distinct cell microvilli, a phenotype not present in colonies lacking Waminoa sp. In most cases, both cnidarian and Acoelomorph hosts displayed high specificity for genetically distinctive Symbiodinium spp. These observations show that the epizoic worms do not acquire their symbionts from the “host” coral.     

Embryogenesis and larval biology of the ahermatypic scleractinian<Emphasis Type="Italic"> Oculina varicosa</Emphasis>

The ivory tree coral Oculina varicosa (Leseur, 1820) is an ahermatypic branching scleractinian that colonizes limestone ledges at depths of 6–100 m along the Atlantic coast of Florida. This paper describes the development of embryos and larvae from shallow-water O. varicosa, collected at 6–8 m depth in July 1999 off Fort Pierce, Florida (27°32.542 N; 79°58.732 W). The effect of temperature on embryogenesis, larval survival, and larval swimming speed were examined in the laboratory. Ontogenetic changes in geotaxis and phototaxis were also investigated. Embryos developed via spiral cleavage from small (100 µm), negatively buoyant eggs. Ciliated larvae developed after 6–9 h at 25°C. Embryogenesis ceased at 10°C, was inhibited at 17°C, and progressed normally at 25°C and 30°C. Larval survival, however, was high across the full range of experimental temperatures (11–31°C), although mortality increased in the warmest treatments (26°C and 31°C). Larval swimming speed was highest at 25°C, and lower at the temperature extremes (5°C and 35°C). An ontogenetic change in geotaxis was observed; newly ciliated larvae swam to the water surface and remained there for approximately 18 h, after which they swam briefly throughout the water column, then became demersal. Early larvae showed no response to light stimulation, but at 14 and 23 days larvae appeared to exhibit negatively phototactic behavior. Although low temperatures inhibited the development of O. varicosa embryos, the larvae survived temperature extremes for extended periods of time. Ontogenetic changes in larval behavior may ensure that competent larvae are close to the benthos to facilitate settlement. Previous experiments on survival, swimming speeds, and observations on behavior of O. varicosa larvae from deep-water adults indicate that there is no difference between larvae of the deep and shallow populations.Communicated by J.P. Grassle, New Brunswick     

Bleaching response of corals and their <Emphasis Type="Italic">Symbiodinium</Emphasis> communities in southern Africa

The high-latitude coral communities of southern Africa suffered minimal impacts during past mass bleaching events. Recent reports indicate an increase in bleaching frequency during the last decade, yet the actual levels of thermal stress and contributing factors in these bleaching events, and the degree of acclimatisation or adaptation on these reefs are poorly understood. During the 2005 warm-water anomaly in the southern Indian Ocean we conducted bleaching surveys and collected samples for genotyping of the algal symbiont communities at 21 sites in southern Mozambique and South Africa. Coral bleaching reached unprecedented levels and was negatively correlated with both latitude and water depths. Stylophora pistillata and Montipora were the most susceptible taxa, whereas three common branching corals had significantly different bleaching responses (Stylophora > Acropora > Pocillopora). Temperature records indicated that localised strong upwelling events coupled with persistent above-average seawater temperatures may result in accumulated thermal stress leading to bleaching. Symbiodinium in 139 scleractinian corals belonged almost exclusively to clade C, with clade D symbionts present in only 3% of the colonies. Two atypical C subclades were present in Stylophora and Pocillopora colonies and these were more abundant in shallow than deeper sites. Taxon-specific differences in bleaching responses were unrelated to different clades of algal symbionts and suggest that Symbiodinium C subtypes with diverse thermal tolerance, coupled with acclimatisation and morphology of the host colony influence the bleaching response. Additionally, the predominance of putatively thermal-sensitive Symbiodinium in southern African corals may reflect a limited experience of bleaching and emphasises the vulnerability of these reefs to moderate levels of thermal stress.     

Photosynthetic characteristics of Prochloron sp./ascidian symbioses

The prokaryotic green alga Prochloron sp. (Prochlorophyta) is found in symbiotic association with colonial didemnid ascidians that inhabit warm tropical waters in a broad range of light environments. We sought to determine the light-adaptation features of this alga in relation to the natural light environments in which the symbioses are found, and to characterize the temperature sensitivity of photosynthesis and respiration of Prochloron sp. in order to assess its physiological role in the productivity and distribution of the symbiosis. Colonies of the host ascidian Lissoclinum patella were collected from exposed and shaded habitats in a shallow lagoon in Palau, West Caroline Islands, during February and March, 1983. Some colonies from the two light habitats were maintained under conditions of high light (2 200 E m^–2 s^–1) and low light (400 E m^–2 s^–1) in running seawater tanks. The environments were characterized in terms of daily light quantum fluxes, daily periods of light-saturated photosynthesis (H_sat), and photon flux density levels. Prochloron sp. cells were isolated from the hosts and examined for their photosynthesis vs irradiance relationships, respiration, pigment content and photosynthetic unit features. In addition, daily P:R ratios, photosynthetic quotients, carbon balances and photosynthetic carbon release were also characterized. It was found that Prochloron sp. cells from low-light colonies possessed lower chlorophyll a/b ratios, larger photosynthetic units sizes based on both reaction I and reaction II, similar numbers of reaction center I and reaction center II per cell, lower respiration levels, and lower P_max values than cells from high-light colonies. Cells isolated from low-light colonies showed photoinhibition of P_max at photon flux densities above 800 E m^–2 s^–1. However, because the host tissue attenuates about 60 to 80% of the incident irradiance, it is unlikely that these cells are normally photoinhibited in hospite. Collectively, the light-adaptation features of Prochloron sp. were more similar to those of eukaryotic algae and vascular plant chloroplasts than to those of cyanobacteria, and the responses were more sensitive to the daily flux of photosynthetic quantum than to photon flux density per se. Calculation of daily minimum carbon balances indicated that, though high-light cells had daily P:R ratios of 1.0 compared to 4.6 for low-light cells, the cells from the two different light environments showed nearly identical daily carbon gains. Cells isolated from high-light colonies released between 15 and 20% of their photosynthetically-fixed carbon, levels sufficient to be important in the nutrition of the host. Q₁₀ responses of photosynthesis and respiration in Prochloron sp. cells exposed briefly (15–45 min) to temperatures between 15° and 45°C revealed a discontinuity in the photosynthetic response at the ambient growth temperatures. The photosynthetic rates were found to be more than twice as sensitive to temperatures below ambient (Q₁₀=3.47) than to temperatures above ambient (Q₁₀=1.47). The Q₁₀ for respiration was constant (Q₁₀=1.66) over the temperature range examined. It appears that the photosynthetic temperature sensitivity of Prochloron sp. may restrict its distribution to warmer tropical waters. The ecological implications of these findings are discussed in relation to published data on other symbiotic systems and free-living algae.     

Embryogenesis and acquisition of algal symbionts by planulae of Xenia umbellata (Octocorallia: Alcyonacea)

Early embryogenesis of the internally brooding soft coral Xenia umbellata and acquisition of algal symbionts in the course of its planular ontogenesis have been examined by scanning and transmission electron microscopy and by light microscopy. The endoderm of adult X. umbellata harbours symbionts mainly in the tentacles and in the peripheral solenia system. The colonies are gonochoric brooders. Algal symbionts were never found in the sperm sacs, and were only rarely found in the follicular tissue enclosing the oocytes. Fertilized eggs pass into endodermal brood pouches where embryogenesis occurs. Cleavage is holoblastic and leads to formation of a solid blastula. Algal symbionts are conspicuously embedded in the parental mesoglea that coats the young embryo, most probably transmitted by surface adherence. At a further stage, this integument disappears and the algae reside extracellularly among the cells of the newly-formed blastula. After subsequent cell proliferation developing planulae possess an inner mass of yolk-laden cells that contain numerous symbiotic algae. Gradually the yolk disintegrates, leaving a cavity enclosed by ectoderm, a thin mesoglea and an inner endoderm with intracellular symbionts. The mature planulae have already been provided with numerous intracellular symbionts by the time they are expelled from the brood pouches. The markedly early symbiont acquisition by the embryos of X. umbellata may help support their developmental requirements in the course of planular ontogenesis.     

Transmission of symbiotic dinoflagellates through the sexual cycle of the host scyphozoan Linuche unguiculata

Intracellular symbiotic dinoflagellates are associated with the tropical scyphozoan Linuche unguiculata (Swartz, 1788) throughout all stages of the host's life cycle. During sexual reproduction, eggs are released in mucus strands that contain symbiotic dinoflagellates. Fertilization and development take place externally in the water column. Epifluorescence and transmission electron microscopy showed that unfertilized eggs did not contain intracellular algae, but that infection of the developing embryo was generally successful by the 128-cell stage (10 h after fertilization at 23° C). However, experiments with artificially provided Cellufluor-labeled algae demonstrated that older embryos and planulae could be infected by algae through at least 24 h post-fertilization, indicating that the L. unguiculata symbiosis represents a semi-closed system. This novel mode of symbiont acquisition results in most sexually-produced offspring becoming infected with maternally-transmitted algae during early development, but allows for acquisition of non-maternally-provided algae later in development. Most of the algal symbionts during the early stages of embryonic and larval development are located within ectodermal cells. This is in contrast to the other life-cycle stages of L. unguiculata (i.e., scyphistoma, medusa, ephyra), where symbionts are found within the gastrodermis of the host.     

Light-enhanced growth of the ascidian Didemnum molle/Prochloron sp. symbiosis

To obtain a direct measurement of the importance of Prochloron sp. to the ascidian Didemnum molle, ascidian colonies from Lizard Island Lagoon, Great Barrier Reef, were grown for 9 d at 0, 10, 40 and 100% sunlight in situ using unidirectional flow chambers. Growth (wet weight) was enhanced up to 40% of full sunlight, at which point growth appears to have been light-saturated. Colonies in 10 and 40% sunlight responded by (1) climbing up the sides of the growth chambers, and (2) flattening out to a more encrusting morphology; also (3) the chlorophyll content of three colonies in zero sunlight decreased by >80%, yet the ascidians remained healthy and did not lose weight. These data show that although the symbiosis may not be obligatory for D. molle, the ascidian's growth is enhanced by Prochloron sp., and the morphology of the ascidian colony is affected by its photobiology.Contribution No. 344 of the Australian Institute of Marine ScienceCommunicated by G. F. Humphrey, Sydney     

Distributions and population structures of two intertidal estuarine polychaetes in the lower St. Lawrence Estuary,with special reference to environmental factors

An intensive study of the spatial distribution ofNereis virens (Sars) andNephtys caeca (Fabricius) was conducted during the spring and autumn of 1986 in the lower St. Lawrence estuary. Statistical analysis showed that spatial variations in density, individual body weight and sexual maturity, particularly in the case ofNereis virens, are correlated with the intertidal level, with certain sediment characteristics, and with the thickness of the colonizable sediment layer. The density ofN. virens increases in an offshore-onshore direction, whereas that ofNephtys caeca decreases in the same direction. For both species, mean body weight increases downshore from the upper intertidal level. Other specific relationships exist in relation to sediment characteristics. Sexually matureNereis virens are found only at the lower intertidal level; sexual maturity inNephtys caeca was not studied. There were no changes in spatial distribution patterns between spring and autumn. The spatial distribution ofNereis virens parallels depth contours and may reflect its ability to inhabit environments which become more physically unstable in an offshore-onshore direction. This spatial distribution is consistent with a model whereby larvae are recruited in the upper intertidal zone and juveniles migrate downshore.     

Effects on larval crabs of exposure to algal toxin via ingestion of heterotrophic prey

We tested whether ingesting toxic algae by heterotrophic prey affected their nutritional value to crab larval predators, using toxic algal strains that are either ingested directly by larval crabs or rejected by them. Ingestion of toxic strains of the dinoflagellates Alexandrium andersoni and A. fundyense by the rotifer Brachionus plicatilis was confirmed. Rotifers having ingested either algal type for five days were fed to freshly hatched larvae of three crab species, with larval survival and stage durations determined. For both algal/rotifer treatments in all three crab species, larvae fed algae directly died during the first zoeal stage, while those fed rotifers that had been fed either algal strain survived to the experiment’s end (zoeal stage 3). Survival was lower, and stage duration longer, for larvae fed rotifers cultured on toxic algae when compared to those fed non-toxic algae. The role of toxic algae in the planktonic food web may be influenced by its direct or indirect ingestion by larval crabs.     

The response of an insect parasitoid,Ormia ochracea (Tachinidae), to the uncertainty of larval success during infestation

Ormia ochracea is a parasitoid fly which lays its larvae on its hosts, the field crickets Gryllus integer and Gryllus rubens, in two distinct modes: (1) directly on the host and (2) around the host. In the field, 12.7% of male crickets were parasitized and 3.2% were super-parasitized. Despite the disadvantages of parasitizing infested hosts, there was no evidence that O. ochracea avoided superparasitism. This and other experiments suggest that the host assessment ability of O. ochracea is less than that reported for many hymenopteran parasitoids. by manipulating the number of larvae in each cricket, we determined that four to five larvae per host resulted in the largest number of adult flies. However, as larval number per host increased from one to six, pupal size, and hence adult size, declined. In the field, hosts were found with a mean of 1.7 ± 1.0 (SD) larvae per cricket, suggesting that there may be some selection pressure against larger clutch sizes. Nevertheless clutch sizes larger than the host can support were sometimes found in the field. During the first mode of larviposition, gravid flies deposited no more than three larvae directly onto the host. Larvae deposited directly on the host had a high probability of infesting it. During the second mode of larviposition, gravid flies laid a larger number of larvae around the host (6.1 ± 5.2). Larvae that were laid around the host were less likely to infest a cricket than were larvae that were deposited directly onto it. O. ochracea is unique in that its two different modes of larviposition have different probabilities of larval success. Even though the success rate for larvae laid during the second mode of larviposition was low, the possibility of parasitizing more hosts appears to have selected for flies laying more larvae (e.g. increasing clutch size) than is optimal if all the larvae successfully entered a single host.     

Mode of zooxanthella transmission does not affect zooxanthella diversity in acroporid corals

Two distinct modes of algal endosymbiont acquisition exist in corals, a direct transmission from the parental colony to the eggs and a larval or post-larval uptake from the environment. The former, maternal-transmission mode is expected to be a more closed system, while the latter is believed to be an open system. Here we test the hypothesis that the diversity of symbionts in closed systems is lower than that in open systems. We examine the identity and diversity of the algal endosymbionts (zooxanthellae) in 25 Montipora species sampled from Irian Jaya (Indonesia) and Magnetic Island (central Great Barrier Reef) and compare the results with those previously obtained from Acropora species, which belong to the same family. All Montipora colonies examined harbour clade C zooxanthellae, with two colonies harbouring both clade C and D zooxanthellae simultaneously. Two algal strains (named C· and D· in this study) present in Montipora have not been observed in Acropora, and may have co-evolved with Montipora. Symbiodinium C· shows approximately 5% sequence divergence from C strains observed in Acropora spp. and occurs in 76% of the colonies examined. Nevertheless, several other C strains commonly found in other corals occur in some of the Montipora colonies. Montipora species transmit their algal endosymbionts directly to the eggs, while Acropora species have to acquire zooxanthellae from the environment every generation. Contrary to our expectations, the diversity of zooxanthellae is similar for the two genera, indicating that the mode of symbiont transmission (i.e. maternal versus horizontal) does not affect symbiont diversity in acroporid corals.Communicated by M.S. Johnson, Crawley     

Furanocoumarins and their detoxification in a tri-trophic interaction

The parsnip webworm, Depressaria pastinacella, specializes on wild parsnip, Pastinaca sativa, and several species of Heracleum, hostplants rich in toxic furanocoumarins. Rates of furanocoumarin metabolism in this species are among the highest known for any insect. Within its native range in Europe, webworms are heavily parasitized by the polyembryonic parasitoid wasp Copidosoma sosares. In this study, we determined whether these parasitoids are exposed to furanocoumarins in host hemolymph, whether they can metabolize furanocoumarins, and whether parasitism influences the ability of webworms to detoxify furanocoumarins. Hemolymph of webworms fed artificial diet containing 0.3 % fresh weight xanthotoxin, a furanocoumarin prevalent in wild parsnip hosts, contained trace amounts of this toxin; as well, hemolymph of webworms consuming P. sativa flowers and fruits contained trace amounts of six of seven furanocoumarins present in the hostplant. Thus, parasitoids likely encounter furanocoumarins in host hemolymph. Assays of xanthotoxin metabolism in C. sosares failed to show any ability to metabolize this compound. Parasitized webworms, collected from populations of Heracleum sphondylium in the Netherlands in 2004, were on average 55 % larger by weight than unparasitized individuals. This weight is inclusive of host and parasitoid masses. Absolute rates of detoxification (nmoles min⁻¹) of five different furanocoumarins were indistinguishable between parasitized and unparasitized ultimate instars, suggesting that the intrinsic rates of metabolism are fixed. Thus, although parasitized larvae are larger, detoxification rates are not commensurate with size; rates in parasitized larvae expressed per gram of larval mass were 25 % lower than in unparasitized larvae.     

Morphological evidence for vertical transmission of symbiotic bacteria in the viviparous sponge<Emphasis Type="Italic"> Halisarca dujardini</Emphasis> Johnston (Porifera,Demospongiae, Halisarcida)

All stages of vertical transmission of symbiotic bacteria, from the penetration into oocytes to the formation of rhagon, were investigated in the White Sea (Arctic) representatives of Halisarca dujardini Johnston (Demospongiae). Small populations of free-living specific symbiotic bacteria inhabit the mesohyl of H. dujardini. They are represented by a single morphotype of small spiral gram-positive bacteria. Vertical transmission of symbiotic bacteria between generations in sponges may occur in different ways. In the case of H. dujardini the bacteria penetrate into growing oocytes by endocytosis. A part of the bacteria plays a trophic role for oocytes and the other part remains undigested in membrane-bound vacuoles within the cytoplasm. In cleaving embryos bacteria are situated between the blastomeres or in the vacuoles. In the blastula all bacteria are disposed in the blastocoel. The symbionts are situated in intercellular spaces in free-swimming larvae and during metamorphosis. Symbiotic bacteria do not play any trophic role in the period of embryonic and postembryonic development of H. dujardini. No signs of destruction and digestion of bacteria were revealed at any stage of development.Communicated by O. Kinne, Oldendorf/Luhe