Experimental induction of gonadal maturation and spawning in the giant jellyfish <Emphasis Type="Italic">Nemopilema nomurai</Emphasis> (Scyphozoa: Rhizostomeae)

We studied the processes of gonadal maturation, spawning, fertilization and embryonic development of the giant jellyfish Nemopilema nomurai, which has been blooming in recent years in the Sea of Japan. Healthy medusae actively swimming offshore had immature gonads, but damaged and ashore-drifted animals had relatively mature gonads. The animals maintained in a small net on the nearshore to mimic the drifted condition showed induced gonadal maturation by the increase in oocyte diameter and darkness in the matrix. A similar maturation process also occurred in isolated pieces of ovary incubated at the same temperature. Fully grown oocytes that probably stopped at the prophase of the first meiosis reinitiated maturation divisions in response to light exposure, and massive spawnings occurred 80–100 min after the exposure. The spawned oocytes were first released within the subgenital sinus, then transported peripherally and finally shed into the gastrovascular cavity. Maturation and spawning in male gonads were similar to those of females except that the male spawning occurred within 30 min after the light exposure and always preceded the female spawning. Hence, it was suggested that fertilization might take place in the female gastrovascular cavity by the sperm that came in from surrounding seawater by animals’ pumping activities after dawn. Having mechanical damages might promote fertilization success of this species in the Sea of Japan.     

Defensive volatile secretions of two diplopod species attract the carrion ball roller scarab <Emphasis Type="Italic">Canthon morsei</Emphasis> (Coleoptera: Scarabaeidae)

Summary. We report field and laboratory evidence indicating that the defensive compounds secreted by the millipedes Amplinus bitumidus (Diplopoda: Polydesmida) and Anadenobolus putealis (Diplopoda: Spirobolida) attract the carrion scarab Canthon morsei (Scarabaeidae: Scarabaeinae). In a Mexican tropical rain forest, C. morsei was mostly caught in pitfall traps baited with the carcasses of both millipede species. Only a few beetles were attracted to fish carrion. Experimental evidence indicates that C. morsei is attracted to freshly dead millipedes. Benzaldehyde and HCN were identified from Amplinus bitumidus; but 2-methyl-1,4-benzoquinone (toluquinone) and 2-methoxy- 3-methyl-1,4-benzoquinone from Anadenobolus putealis. Benzaldehyde attracted significantly more C. morsei individuals than fish meat, or millipedes of both species did after rinsing in hexane. This is the first report of Polydesmida attracting dung beetles.     

Turnover and vertical transport of zinc by the euphausiid Meganyctiphanes norvegica in the Ligurian Sea

Participatory turnover time is defined as the time required to cycle an element in a system through a given material in that system. The participatory turnover time of ionic zinc by the adult Meganyctiphanes norvegica population in the Ligurian Sea ranged between 498 and 1243 years, depending upon the available food supply, and considering the food chain as the only route for zinc accumulation by the population. A total-impact turnover time was calculated as the sum of the participatory turnover time for live individuals plus the time required for dead euphausiids to lose 90% of their zinc to the water. Carcasses lost zinc to the water slower than either feces or molts, and so established the maximum loss time for all particulate excretion products; nevertheless, total-impact turnover time for zinc did not differ significantly from the participatory turnover time. The net vertical transport of zinc by M. norvegica from the sea surface to any specified depth can be calculated as the sum of the dissolved zinc excreted below the depth plus the concentrations of zinc left in feces, molts, and carcasses after they have sunk to the specified depth. Carcasses sink the fastest and lose the smallest fraction of their zinc concentration during descent; fecal pellets sink the slowest and lose the greatest fraction of their zinc concentration, and molts are intermediate. Nevertheless, feces represents the major route for delivering zinc to the bottom of the Ligurian Sea (2500 m), because concentration of the element in the pellets is so much higher than in carcasses or molts. Excretion of dissolved zinc into the water at the vertical migration depth of the living population during daylight hours was also inconsequential. Feces zinc represented over 80% of the total zinc transported to the sea floor if only marginal food supplies were available to the euphausiids, and over 90% if food was in sufficient supply. M. norvegica can effect a net transport of about 98% of its body zinc concentration below 500 m daily, in conditions of sufficient food supply and assuming that no released products are eaten during descent. If the food supply in the Ligurian Sea is considered only marginal throughout the year, M. norvegica can still effect a daily net transport below 500 m of about 36% of its body concentration, and about 6% of its body concentration will reach 2500 m daily.     

Seasonal patterns in size and abundance of <Emphasis Type="Italic">Phyllorhiza punctata</Emphasis>: an invasive scyphomedusa in coastal Georgia (USA)

Phyllorhiza punctata, commonly called the Australian white spotted jellyfish, invaded the Caribbean in the 1960s, becoming established there and subsequently in the United States in the northern Gulf of Mexico (by 2000) and eastern Florida (2001). With the prevailing Loop Current flowing clockwise around the Gulf of Mexico and joining the Gulf Stream along eastern Florida, potential transport of P. punctata along the eastern seaboard of the USA could be facilitated. P. punctata medusae were collected in small numbers along the entire Georgia coast during May–November in 2007 and 2008. Medusa bell diameters increased both years from ca. 10 cm in May to ca. 33 cm in autumn. Specimens lacked zooxanthellae, as reported for medusae in the northern Gulf of Mexico and Florida. It is possible that the P. punctata medusae observed were transported from established populations to the south; however, whether or not this species is established along the Georgia coast has yet to be determined.     

Invading <Emphasis Type="Italic">Aurelia aurita</Emphasis> has established scyphistoma populations in the Caspian Sea

The Caspian Sea has no endemic Scyphozoa. In 1999, a mass accumulation of Aurelia medusae was recorded, indicating that sometime earlier, jellyfish had invaded the basin, but since then no scyphozoans have been reported in the Caspian. In the fall of 2008, we found scyphistomae (scyphoid polyps) during a cruise to the eastern Middle Caspian. The scyphistomae were numerous (100–10,000 ind. m⁻²) and occupied a depth range of 30–73 m. Genetic data (18S rDNA, ITS-1 and COI) showed that the scyphistomae belonged to the species A. aurita. The current genetic data set is insufficient to determine the source region(s) of the invasive A. aurita. It remains unclear why no moon jellies have been recorded in the Caspian in the last 10 years. Because swarming scyphomedusae are often pests, the presence of scyphistomae should be considered as a warning of a possible outbreak of A. aurita medusae in the Caspian.     

Enhanced survival of 0-group gadoid fish under jellyfish umbrellas

Young (0-group) gadoid fish, which have been observed sheltering beneath jellyfish (Scyphozoa and Hydrozoa) umbrellas, may find refugia from predation by retreating among medusan tentacles. The survival of juvenile gadoids may therefore be improved by high abundances of medusae. Jellyfish (including Cyanea lamarckii and C. capillata) were caught in the North Sea during routine sampling for 0-group gadoids (cod Gadus morhua, haddock Melanogrammus aeglefinus, Norway pout Trisopterus esmarkii and whiting Merlangius merlangus) between 1971 and 1986, and considerable overlaps have been shown between the spatial distributions of these fish and medusae. Here correlations are made between the abundance of medusae in the North Sea and the residual survival of 0-group fish, as quantified by the deviation in 1-group recruitment from the expected Ricker modelled estimate. Significant positive correlations between the residual survival of whiting and medusa abundance are evident for each individual Cyanea sp. and for the combined Cyanea spp. ln(maximum) abundance (all R ≥ 0.60, P < 0.01, N = 15). The abundance of jellyfish may thus be an important factor influencing the mortality of whiting in the North Sea, and as such should be considered in the development of ‘ecosystem-based’ management of whiting stocks.     

Reproductive biology of the edible jellyfish Catostylus mosaicus (Rhizostomeae)

 The reproductive biology of the jellyfish Catostylus mosaicus (Quoy and Gaimard 1824) (Scyphozoa: Rhizostomeae) was investigated in New South Wales, Australia. Medusae were gonochoristic. There was a 1:1 ratio of male and female medusae and there was no evidence of sexual dimorphism. Oocytes arose from the gastrodermis and maintained contact with the gastrodermis, via trophocytes, throughout gametogenesis. Spermatogenesis occurred within follicles that arose from invagination of the gastrodermis. Detailed sampling of gonads over a period of 3.25 yr in Botany Bay, and over 2.5 yr in Lake Illawarra, indicated that gametogenesis occurred almost continuously during the year. Oocytes were smaller, or were absent from the ovaries during 3 of the 4 winters sampled at Botany Bay and during all 3 winter periods sampled at Lake Illawarra. Comparisons were made with other locations, although these were sampled less frequently. When medusae were present at a location, similar trends were observed. The size at which medusae matured varied, but during non-winter periods and at two locations, all medusae exceeding 130 mm diam were considered mature. Received: 6 January 2000 / Accepted: 3 July 2000     

Further studies on the pressure tolerance of deep-sea crustacea,with observations using a new high-pressure trap

A deep-sea benthic trap is described with which amphipods (Tmetonyx cicada) were both collected and observed at their ambient pressure of 134 atm. T. cicada collected with decompression from the same depth and locality were more sensitive to subsequent recompression than those amphipods brought to the surface at their ambient pressure. T. cicada from 2700 m experience irreversible injury during decompression to atmospheric pressure. The pressure tolerance of the deep-sea mysid Gnathophausia zoea was measured and compared with the tolerance of mid-water decapods and the shallow-water Crangon crangon. G. zoea became more sensitive to high pressure with prolonged exposure to atmospheric pressure. Deep sea animals exhibit a tolerance to high pressure related to their normal ambient pressure; sensitivity to decompression is also related to normal ambient pressure.     

Daily photosynthesis,respiration, and carbon budgets in a tropical marine jellyfish (Mastigias sp.)

From measured diel photosynthesis and respiration rates, using oxygen electrodes, estimates of carbon flux between symbiotic algae (zooxanthellae) and host animal are presented for the marine scyphomedusan Mastigias sp. from a marine lake in Palau, Western Caroline Islands, during February and March 1982. The carbon budgets calculated for these lake medusae indicate that carbon fixed photosynthetically by zooxanthellae and made available to the host may satisfy up to 100% of the host's daily metabolic carbon demand (CZAR). The stable carbon isotope (¹³C) signature of the mesogleal carbon of lake Mastigias sp. was close to that of the zooxanthellae, supporting the interpretation that while these medusae may feed holozoically, some of their carbon comes from their symbionts. The diel photosynthesis, respiration, and preliminary estimates of carbon budgets of three individuals of another ecotype of Mastigias sp. collected from nearby oceanic lagoons are also given. Photosynthesis of lagoon medusae was generally greater than that for lake medusae of similar size, and lagoon medusae were phototrophic with respect to carbon, with commensurately greater CZAR values. Carbon translocated from the symbiotic algae also may contribute to the growth requirements of both lake and lagoon medusae. From carbon flux data, the lake jellyfish were estimated to contribute about 16% to the total primary productivity of their marine lake habitat.     

Abundance,distribution and prey composition of scyphomedusae in the southern North Sea

The annual cycle of abundance and distribution of the scyphozoan medusae Aurelia aurita, Cyanea lamarckii, C. capillata and Chrysaora hysoscella were studied in the southern North Sea in 2004 and 2005. Three different patterns of seasonal occurrence of medusae were distinguished: (1) the early occurring C. lamarckii (February–August), (2) C. capillata and A. aurita (April–August) and (3) the late appearing C. hysoscella (July/August–September). Cyanea lamarckii was the most frequently encountered species in this study; its highest mean abundance was 1.8 ± 2.7 ind. 100 m⁻³. The prey spectra of C. lamarckii, C. capillata and C. hysoscella contained several copepod and other crustacean species and thus make them potential competitors with fish larvae. Medusae in this study also consumed fish eggs and larvae, including clupeids, in all months analysed. Although peak spawning of sprat (Sprattus sprattus) coincides with the maximum abundance of medusae (May–June) the relative low abundance of all medusae species in this study makes jellyfish predation unlikely to be a factor controlling sprat recruitment in the time frame investigated.     

Chemically mediated competition between microbes and animals: microbes as consumers in food webs

Microbes are known to affect ecosystems and communities as decomposers, pathogens, and mutualists. However, they also may function as classic consumers and competitors with animals if they chemically deter larger consumers from using rich food-falls such as carrion, fruits, and seeds that can represent critical windfalls to both microbes and animals. Microbes often use chemicals (i.e., antibiotics) to compete against other microbes. Thus using chemicals against larger competitors might be expected and could redirect significant energy subsidies from upper trophic levels to the detrital pathway. When we baited traps in a coastal marine ecosystem with fresh vs. microbe-laden fish carrion, fresh carrion attracted 2.6 times as many animals per trap as microbe-laden carrion. This resulted from fresh carrion being found more frequently and from attracting more animals when found. Microbe-laden carrion was four times more likely to be uncolonized by large consumers than was fresh carrion. In the lab, the most common animal found in our traps (the stone crab Menippe mercenaria) ate fresh carrion 2.4 times more frequently than microbe-laden carrion. Bacteria-removal experiments and feeding bioassays using organic extracts of microbe-laden carrion showed that bacteria produced noxious chemicals that deterred animal consumers. Thus bacteria compete with large animal scavengers by rendering carcasses chemically repugnant. Because food-fall resources such as carrion are major food subsidies in many ecosystems, chemically mediated competition between microbes and animals could be an important, common, but underappreciated interaction within many communities.     

The symbiotic relationship between Lychnorhiza lucerna (Scyphozoa, Rhizostomeae) and Libinia spinosa (Decapoda, Epialtidae) in the Río de la Plata (Argentina–Uruguay)

Herein, we characterize a symbiotic relationship between the scyphomedusa Lychnorhiza lucerna and the decapod crustacean Libinia spinosa in Río de la Plata, South America. Of 843 specimens of L. lucerna examined during the study, 69 (8.2 %) hosted L. spinosa within subgenital spaces. The broad spatial and temporal scale of the study, together with the large number of observations made, confirm an association between the two species. Medusae having crab associates were mature and larger than those lacking such symbionts. Adult crabs of both sexes, as well as juveniles and soft-shell individuals, were found as associates of medusae. Analysis of crab stomach contents revealed the presence of nematocysts and copepod remains. Our results suggest that medusae provide protection and possibly access to food for crabs. Benefits related to transportation were not clear and need further evaluation. Crabs of L. spinosa may acquire their scyphozoan symbionts either as larvae planktonic stages or as adult crabs attaching to jellyfish when aggregating close to bottom.     

Occurrence and public perception of jellyfish along the German Baltic coastline

Jellyfish accumulations are often problematic for tourism and fisheries both along European coastlines and overseas. They damage the reputation of several seaside resorts and cause serious economic losses. On the German Baltic coast, jellyfish accumulations are well known, too. Besides population increase due to anthropogenic effects, normal hydrodynamic processes are notable key factors when it comes to jellyfish accumulations. Along German Baltic coastlines, moon jellyfish (Aurelia aurita) occur most frequently but also stinging lion??s mane jellyfish (Cyanea capillata) appear regularly, however not as abundant. The objectives of our study were a) to screen when, where, how many and which jellyfish species occur on German Baltic coasts, b) to find the causes for jellyfish accumulations and c) to evaluate how beach visitors perceive native medusae and whether information can influence their perception. Different methods where combined: systematic jellyfish observations in cooperation with lifeguards, investigations of jellyfish abundance in shallow water zones, and interviews with beach visitors. Our results suggest that jellyfish occurrence along the German Baltic coast correlates with offshore wind or shore-parallel wind, which cause upwelling events. In contrast, there is no evidence that frequency of jellyfish occurrence was dependent on water temperature in summer. In regard to tourism, we found that beach visitors who received information about jellyfish stated to feel significantly less bothered by medusae. Overall, this article addresses different methods to learn more about jellyfish accumulations and it shows information strategies in terms of improved beach management.     

Ciliated epibionts on the Arctic sympagic amphipod<Emphasis Type="Italic"> Gammarus wilkitzkii</Emphasis> as indicators for sympago–benthic coupling

In September 2003 specimens of the sympagic amphipod Gammarus wilkitzkii were sampled in drifting pack ice above 50 m water depth and in the pelagic/benthic environment beneath in the coastal waters off Svalbard (Norway). Abundance values ranged between 1.5 and 8 individuals m^–2 for the pelagic/benthic and the sympagic environments, respectively, and showed corresponding sex ratios of 1:1.4, favoring males. In the ice a significantly higher number of juveniles prevailed. In contrast, ovigerous females were more abundant among pelagic/benthic living specimens. Approximately 25% of the individuals hosted ciliated epibionts of the genera Ephelota, Cryptacineta, Acineta, Podophrya (all suctoria), and Epistylis (peritrichia). Cryptacineta and Ephelota were the most abundant epibionts on this amphipod species. Female specimens of G. wilkitzkii showed the highest degree of infestation (>2,100 individual epibionts per amphipod specimen: indE/A) followed by juveniles (>1,200 indE/A) and males (>220 indE/A). Highest densities of epibionts were found on anterior body parts with the antennae bearing up to 130 individuals. This is the first sighting of epibionts on crustacea from the sympagic environment. We interpret them as biomarkers that indicate the existence of sympago–benthic coupling processes between the ice and the underlying waters and the seafloor. The population structure and the proportion of infested specimens are equal for the amphipods sampled from both the sympagic and the pelagic/benthic environment, indicating the existence of exchange (coupling) processes between the two habitats. Considering the annual ice cycle, local ice drift patterns, and the shallow water depth in the sampling area, we hypothesize that ice amphipods spend ice-free periods near the seafloor where they may serve as basibionts for protozoans and eventually re-colonize the ice with the onset of ice formation. Our observations strongly emphasize that shallow coastal areas serve as both retention (rather than sink) areas for ice fauna during ice-free periods, and as stepping-stones for re-colonizing the ice when the ice is formed in winter.Communicated by M. Kühl, Helsingor     

Life cycle of the rhizostome jellyfish <Emphasis Type="Italic">Rhizostoma octopus</Emphasis> (L.) (Scyphozoa,Rhizostomeae), with studies on cnidocysts and statoliths

Stages in the life cycle of the rhizostome jellyfish Rhizostoma octopus (L.) were reared in the laboratory from planula to young medusa and are described here. Fertilization of eggs was external, and planulae, 110–150 μm in length, appeared after 2 days. These settled and developed into the scyphistoma stage. Scyphistomae reached a maximum size of 2.3 mm, bore up to 24 tentacles, and had a large manubrium. Polyps reproduced asexually mainly by podocysts. Strobilation was induced by temperature change. Each strobila released up to five ephyrae that measured 2.7–5.8 mm in diameter at liberation. During transformation from newly released ephyra to young medusa, velar lappets appeared and increased in number, the manubrium developed eight branched oral arms with epaulettes, and a marginal gastric network arose. Only one cnidocyst type, the “a”-atrichous haploneme, was present in the planula. In addition to these haplonemes, heterotrichous microbasic euryteles were observed in polyps, ephyrae, and medusae. Statoliths, located in the rhopalia, had a characteristic compact shape. These sensory structures increased in number and size with the growth of ephyrae and medusae.     

Rate-temperature responses in scyphozoan medusae and polyps

The effects of temperature on oxygen consumption and spontaneous rhythmic activity have been investigated in various stages of the life histories of 3 species of jellyfish from the Chesapeake Bay, USA. All 3 species clearly show the ability to acclimate positively to temperature change. Thermal sensitivity of metabolism in the winter medusa Cyanea capillata fulva is fairly low at temperature intervals which are experienced in nature. Polyps of the two summer medusae, Chrysaora quinquecirrha and Aurelia aurita, show reduced metabolic sensitivity at temperatures normally accompanying high developmental activity and the onset of strobilation.     

Microphytobenthos of the Dogger Bank: a comparison between shallow and deep areas using phytopigment composition of the sediment

The surface sediment characteristics related to benthic microalgae primary production were studied at the Dogger Bank, North Sea, in order to evaluate the potential role of microphytobenthos as a food source for the macrobenthic fauna. Twenty-one stations were sampled in July 2001 and May 2002, with water depth ranging from 16.3 to 68.5 m. High-performance liquid chromatography pigment analyses revealed that concentrations of chlorophyll a, chlorophyll c and fucoxanthin are mainly associated with benthic diatom flora at most parts of the Dogger Bank. High percentage of phytopigments (>50%) was firmly attached to sand grains at the stations shallower than 40 m water depth. The deeper stations were characterized by a phytopigment composition originating from pelagic phytoplankton settled on the sea floor. Qualitative microscopy showed that the benthic microflora on top of the Bank mainly consists of small diatoms (5–10 μm), such as e.g., Diploneis spp., living attached to the sand grains. The results are discussed concerning possible implications for ecology and biogeochemistry of the Dogger Bank area.     

Life cycle of the jellyfish <Emphasis Type="Italic">Rhizostoma pulmo</Emphasis> (Scyphozoa: Rhizostomeae) and its distribution,seasonality and inter-annual variability along the Catalan coast and the Mar Menor (Spain,NW Mediterranean)

Rhizostoma pulmo is one of the most abundant scyphomedusae along the Mediterranean coasts. To understand changes in the population densities of the medusa stage and its relationship with the benthic stage, we describe all developmental stages from the life cycle of R. pulmo, from the scyphistoma stage to young medusae reared in the laboratory. Mature scyphistomae showed a mean total body length of 1.7 mm and the average tentacle number was 16. Asexual reproduction occurred by lateral budding, podocyst and pedalocyst production. Each strobila released up to 8 ephyrae that had a mean total body diameter of 3.5 mm. Moreover, we also present data on the temporal and spatial distribution of the species in the Catalan Sea and the coastal lagoon on Mar Menor (NW Mediterranean) during the years 2007–2009. In the Catalan Sea, the presence of adult R. pulmo was recorded as extended along the whole coast, but most of the observation was concentrated in the central area; the highest abundances were recorded during the months of July and August. The highest number of observations was detected in 2008 in coincidence with the inter-annual variation observed in the Mar Menor lagoon.