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     

Laboratory observations on the life history of Rhopilema verrilli (Scyphozoa: Rhizostomeae)

The life history of the scyphozoan Rhopilema verrilli is described from the planula to the young medusa stages. Planulae are retained within the gonadal tissue of the medusa until fully developed. On liberation, most planulae set and developed into small scyphistomae within 7 to 10 days. The scyphistoma differs from those of other described species in having an unusually large, clavate manubrium. The only means of asexual reproduction observed in the scyphistoma cultures involved the formation of podocysts. Strobilation was usually of the monodisc variety, although polydics strobilation was not infrequent. The process of strobilation was completed within 7 days at 20°C. Newly liberated ephyrae typically had 8 pairs of lappets, and 8 rhopalia. Ephyral development resembled that of the closely related rhizostome Rhizostoma pulmo. The cnidome of the planula and scyphistoma consisted of atrichous isorhizas (a atrichs) and microbasic heterotrichous euryteles, while that of the strobila and ephyra consisted of a atrichs, euryteles, and holotrichous haplonemesContribution No. 546 from the Virginia Institute of Marine Science, Gloucester Point, Virginia 23062, USA.     

Life cycle of the jellyfish <Emphasis Type="Italic">Lychnorhiza lucerna</Emphasis> (Scyphozoa: Rhizostomeae)

The life cycle of Lychnorhiza lucerna (Scyphozoa: Rhizostomeae) and the settlement preferences of its larvae were studied using laboratory-based rearing experiments. Mature medusae of L. lucerna were collected from the beach of the Río de la Plata estuary, Argentina. This species displayed the typical metagenetic, (i.e. medusoid and polypoid), life cycle reported for other rhizostomes. The fertilized eggs developed into motile and short lived planulae. The majority of planulae settled on the air-water interface (p < 0.001). Of those that settled on the settlement plates provided, no significant differences were observed between styrene slides, glass slides and shells of the bivalve Mactra isabelleana (p > 0.05). No planulae settled on stones. Several hours after planulae settled, they metamorphosed into sessile four-tentacled scyphistomae. Most scyphistomae attached onto the air-water interface. At 19–22°C, the scyphistomae grew up to 22 tentacles and reached 1,500 μm height. The scyphistomae increased their numbers by means of formation of podocysts from which new polyps emerged and strobilated. Strobilation occurred 46 days after settlement. Only polydisk strobilation was observed and each strobila always produced three ephyrae. After releasing ephyrae, strobilae returned to normal scyphistomae and were capable of repeating strobilation. A single founder polyp was estimated to produce up to 60 ephyrae over 4 months. Ephyrae developed into metephyrae 15 days after release at 19–22°C. In this paper we describe the morphological and some behavioural features of L. lucerna in the polypoid and early medusoid stages.     

Recent strandings of the giant jellyfish Rhizostoma luteum Quoy and Gaimard, 1827 (Cnidaria: Scyphozoa: Rhizostomeae) on the Atlantic and Mediterranean coasts

We present reports of sightings of living and stranded specimens of Rhizostoma luteum on the Atlantic coast of Morocco and along the south shore of the Iberian Peninsula in June–July 2012 and in January–February 2013. During summer 2012 and following the dominant currents, the jellyfish first appeared in the Gulf of Cadiz west of the Strait of Gibraltar. Subsequently, seven additional sightings were reported east of the Strait, in the Alboran Sea. In winter 2013, another event of stranded individuals of this species occurred in the Gulf of Cadiz. A phylogenetic analysis performed on the mitochondrial cytochrome c oxidase I (COI) gene sequence in specimens from both stranding events confirmed the morphological classification, ratifying that R. luteum differs from Rhizostoma octopus and Rhizostoma pulmo. This study records the presence of this species for the first time in 60 years.     

Spatial aggregations of the swarming jellyfish Pelagia noctiluca (Scyphozoa)

In the Adriatic Sea, the spatial distribution and aggregation aspects of swarning Pelagia noctiluca (Forskål, 1775) were studied by SCUBA divers from August 1984 to November 1985. Medusae were asually distributed in the upper 20- to 30-m layer. Dense aggregations caused by wind, currents and tidal phenomena occurred only in shallow coastal waters. Maximum population densities of swarms drifting freely near the shore were estimated at about 20 individuals per cubic meter, but when they had drifted ashore jellyfish could reach concentrations of 150 to 600 individuals in the same volume of sea water. Some peculiar forms of aggregations could be distinguished: couples, clusters, and surface and bottom aggregations. The effects of wind, currents and geomorphological features of the shore on passive aggregations of jellyfish are presented.     

Substrate choice and settlement preferences of planula larvae of five Scyphozoa (Cnidaria) from German Bight,North Sea

The settlement behaviour of planula larvae and their development to young polyps was investigated in laboratory experiments in five scyphozoan species [Aurelia aurita (L.), Cyanea capillata (L.), Cyanea lamarckii Péron and Leseur, Chrysaora hysoscella (L.), and Rhizostoma octopus (L.)]. The undersides of settling plates were strongly preferred for settlement. Shells, the only natural substrate type offered, were less attractive than artificial substrates (concrete, machined wood, polyethylene, and glass). The advantages of colonization of substrate undersides for survival and reproduction of polyps are discussed. It is supposed that the increase of artificial substrates in our seas, due to marine litter pollution and submarine building activities, enlarge the areas of distribution of scyphozoan polyps, in coastal as well as in off-shore regions. Subsequent increases in ephyra production by polyps are probably one reason for the increase in mass occurrences of jellyfish recognized worldwide during the last few decades. It is suggested that the early developmental stages in the cnidarian life cycle, the planula larvae, and the polyps, play the key role in the development of jellyfish outbursts.     

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.     

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.     

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.     

Geographic variation and behavioral evolution in marine plankton: the case of<Emphasis Type="Italic"> Mastigias</Emphasis> (Scyphozoa,Rhizostomeae)

Although complex behavior in marine zooplankton has been considered strong evidence of adaptation, ethological studies of marine zooplankton generally have not employed either the comparative approach or evolutionary perspective necessary to distinguish adaptation from any alternative. Consequently, the potential for intra-specific variation in the behavior of marine zooplankton has received insufficient attention and conclusions of adaptation remain poorly substantiated. Intra-specific comparison of patterns of migration and behavior for seven populations of golden jellyfish, Mastigias (Scyphozoa: Rhizostomeae), inhabiting isolated marine lakes and semi-enclosed lagoon coves in Palau document population specific differences in patterns of horizontal migration, vertical migration, pulse rate, swimming speed, and turning behavior. Evidence was found for symplesiomorphic behaviors, canalization, exaptation, adaptation, and probably once-deleterious traits. Behavioral evolution likely proceeded via, at least, relaxation of selection, trade-offs with morphology, and natural selection effected by predation. Behavioral patterns also may change with ontogeny. Geographic variation in the behavior of marine plankton therefore can be substantial and patterns of evolution complex. Behavioral evolution can rapidly generate coastal biodiversity. Thus, geographic variation in marine plankton is of potential interest to ethologists, evolutionary biologists, biogeographers, and conservation biologists.Electronic Supplementary Supplementary material is available in the online version of this article at Communicated by G.F. Humphrey, Sydney     

Global phylogeography of<Emphasis Type="Italic"> Cassiopea</Emphasis> (Scyphozoa: Rhizostomeae): molecular evidence for cryptic species and multiple invasions of the Hawaiian Islands

The upside-down jellyfish Cassiopea is a globally distributed, semi-sessile, planktonically dispersed scyphomedusa. Cassiopea occurs in shallow, tropical inshore marine waters on sandy mudflats and is generally associated with mangrove-dominated habitats. Controversy over the taxonomy of upside-down jellyfishes precedes their introduction to the Hawaiian Islands during the Second World War, and persists today. Here we address the global phylogeography and molecular systematics of the three currently recognized species: Cassiopea andromeda, C. frondosa, and C. xamachana. Mitochondrial cytochrome c oxidase I (COI) sequences from Australia, Bermuda, Fiji, the Florida Keys, the Hawaiian Islands, Indonesia, Palau, Panama, Papua New Guinea, and the Red Sea were analyzed. Highly divergent COI haplotypes within the putative species C. andromeda (23.4% Kimura 2-parameter molecular divergence), and shared haplotypes among populations of two separate putative species, C. andromeda and C. xamachana from different ocean basins, suggest multiple anthropogenic introductions and systematic confusion. Two deeply divergent Oahu haplotypes (20.3%) from morphologically similar, geographically separate invasive populations indicate long-term (14–40 million years ago) reproductive isolation of phylogenetically distinct source populations and cryptic species. Data support at least two independent introductions to the Hawaiian Islands, one from the Indo-Pacific, another from the western Atlantic/Red Sea. Molecular phylogenetic results support six species: (1) C. frondosa, western Atlantic (2) C. andromeda, Red Sea/western Atlantic/Hawaiian Islands (3) C. ornata, Indonesia/Palau/Fiji (4) Cassiopea sp. 1, eastern Australia (5) Cassiopea sp. 2, Papua New Guinea and (6) Cassiopea sp. 3, Papua New Guinea/Hawaiian Islands.Communicated by P.W. Sammarco, Chauvin     

Bioluminescence of deep-sea coronate medusae (Cnidaria: Scyphozoa)

Bioluminescence is the production of visible light by a living organism. The light commonly appears as flashes from point sources (involving one or more cells, usually described as photocytes) or as a glandular secretion. A visible flash usually involves synchronous light emission from a group of cells or, if from a single-celled organism such as a dinoflagellate, from a group of organelles. The number of cells (or organelles) responding synchronously is the main determinant of the flash intensity. Bioluminescence is a common phenomenon in many deep-sea animals and is widespread among the Cnidaria. In this paper, we compare and contrast in situ and laboratory recordings of the bioluminescent responses of specimens of the deep-sea scyphozoans Atolla wyvillei, Atolla vanhoffeni, Atolla parva, Nausithoe rubra, Paraphyllina intermedia, Periphyllopsis braueri and Periphylla periphylla. Displays in all seven species consist of localised flashes and propagated waves of light in the surface epithelium. The first few single waves propagate at rates of up to 60 cm s^-1 but subsequent ones in any sequence of stimuli gradually decrease in speed. After several single wave responses, a subsequent stimulus may elicit multiple waves that persist for several seconds. Following such a frenzy, the specimen becomes temporarily refractory to further stimuli, but if rested will recover its normal responses and may produce further frenzies. The dome area, situated above the coronal groove, of the genera Paraphyllina, Periphylla, and Nausithoe is covered with luminescent point sources. Such point sources are generally absent from the dome of species of Atolla. Captured specimens of A. parva also produce secretory bioluminescence, corroborating prior in situ observations of this ability. Secretory bioluminescence in P. periphylla takes the form of scintillating particles released from the lappet margins. We did not observe secretory displays in specimens of any other species in the laboratory, but one instance of apparent secretory luminescence was recorded in situ in a specimen of A. wyvillei.Communicated by J. P. Thorpe, Port Erin     

Life history of the sandy-beach amphipod Neohaustorius schmitzi (Crustacea: Haustoriidae)

Neohaustorius schmitzi Bousfield, the dominant macroscopic invertebrate on sandy beaches in North Carolina, USA, is strictly intertidal, and distributed from mean low tide to high tide, with an average density of 790/m². Location of greatest densities within the intertidal zone is dependent upon season and sex. An aggregated distribution is shown. N. schmitzi has 2 generations/year: a winter generation which lives about 8 months, and a summer generation which lives about 4 months. The reproductive season extends from February through October, with spring and summer peaks. Each female appears to produce only 1 brood of young. The number of young ranges from 2 to 14, depending upon the length and generation of the female.     

Life history and population dynamics ofKinbergonuphis simoni (Polychaeta: Onuphidae)

The relationship between life-history characteristics and population dynamics were investigated in the onuphid polychaeteKinbergonuphis simoni (Santos, Day and Rice) between 1982 and 1987. The studied population is located in Upper Tampa Bay, Florida, USA. This worm attains sexual maturity at 5 to 10 mo of age, depending on temperature. Several consecutive broods are produced during a female's life time, with 7 to 26 young per brood. Adults die in June–July after the breeding season. Generations of breeding individuals do not overlap and the replacement of generations occurs in July–August. Only one extended breeding season is experienced per life time. Life span does not exceed 2 yr. Field population density in 1982 was high in fall and spring (2000 to 9000 individuals/m²), low in summer (1600 individuals/m²), and slightly depressed in winter (2500 to 4000 individuals/m²). Changes in population density may be explained by seasonal, temperature-controlled changes in instantaneous birth and death rates. Birth rates are high in fall and spring, while death rates are high in early summer. Death of juveniles contributes to the winter density decrease.     

Morphology and development of benthic and pelagic life stages of North Sea jellyfish (Scyphozoa, Cnidaria) with special emphasis on the identification of ephyra stages

Jellyfish blooms or invasions could be detected in an early phase of development if the youngest medusa stages (ephyrae) and their early growth stages (post-ephyrae) were identifiable in plankton samples but a useful identification key for ephyrae in early growth stages is lacking for most species. In the present study, the identification characteristics of adult North Sea scyphomedusae (Aurelia aurita, Cyanea capillata, Cyanea lamarckii, Chrysaora hysoscella, Rhizostoma octopus) collected around the island of Helgoland (German Bight) in July?CAugust 2003 and 2004 are described. Planula larvae were measured and reared to polyps in the laboratory. The process of ephyrae development asexually produced by the polyps (strobilation) was photo-documented. Photographs of the ephyrae growth stages were combined with drawings of features useful for the species identification. The provided identification key allows discrimination among post-ephyrae from plankton samples, probably leading to conclusions on the development of jellyfish blooms and their causes.     

Some carotenoids in the jelly-fish Aurelia aurita (Scyphozoa: Discomedusae)

The author investigated the carotenoids in the jelly-fish Aurelia aurita (L.), from the Baltic Sea. By means of columnar and thin-layer chromatography, the following carotenoids were found to be present in that species: -carotene, echinenone isocryptoxanthin, canthaxanthin, lulein, zeaxanthin, astaxanthin and astacene.     

Life history and fishery of Lepidopus caudatus (Pisces: Trichiuridae) in the Catalan Sea (Northwestern Mediterranean)

The frostfish Lepidopus caudatus (Euphrasen, 1788) is a mesopelagic species; it occurs along the shelf and slope down to 450 m in the Catalan Sea. Its' fishery, population structure, growth, diet and reproduction was studied on the basis of commercial statistics from 1984 to 1990 and on samplings made in 1988 and 1989. L. caudatus is fished by two types of commercial method, trawl and long-line; catches by the latter method display a marked seasonality. The minimum size of fish caught is 26 cm total length by trawling and 71 cm by long-line. Maximum sizes are similar for both gears: 196 and 188 cm, respectively. L. caudatus exhibits fast growth and attains an age of 8 yr. It feeds on small mesopelagic prey. It is a partial spawner reproductive activity occurring from April to November. Males attain sexual maturity at 97 cm and females at 111 cm total length.