Population dynamics of natural colonies of <Emphasis Type="Italic">Aurelia</Emphasis> sp. scyphistomae in Tasmania,Australia

The aim of this study was to identify potential environmental controls of the asexual phases of reproduction by measuring the rates of asexual reproduction (budding and strobilation) and mortality in naturally occurring populations of Aurelia sp. scyphistomae at different spatial and temporal scales. The percentage cover, density of colonies of Aurelia sp. scyphistomae, and density of the population of two naturally occurring colonies of Aurelia sp. scyphistomae were examined over 2 years in southern Tasmania. Artificial substrates were also deployed to investigate colony dynamics when density dependent effects were reduced. Clear spatial and temporal differences in the population dynamics of the colonies were observed. Density dependent effects controlled budding and recruitment of new scyphistomae to the substrate when populations were dense and space limiting. In contrast, environmental controls of budding and strobilation were more apparent in a colony with significantly greater area of bare substrate and hence room for expansion. Water temperature and rainfall (as a proxy for salinity) were linked to changes in population size. Annual and seasonal differences in population dynamics were not observed in a colony limited by space but were apparent in a colony where space was not limited. When space was removed as a limiting factor by deploying artificial substrates, a seasonal environmental effect on the rate of growth of the colony was observed. These studies suggest that the growth, survival and reproduction of the sessile colonial phase of Aurelia sp. is regulated by a combination of density dependent factors and environmental conditions, which are consequently important to the formation of jellyfish blooms.     

Allozyme and morphological evidence for a newly introduced species of Aurelia in San Francisco Bay,California

The jellyfish Aurelia aurita (Linnaeus, 1758) is usually considered to be a cosmopolitan species. Aurelia sp. medusae observed at Foster City, San Francisco Bay, California, USA, since 1988 are morphologically distinct from Aurelia sp. collected 200 km away in Monterey Bay, but are morphologically similar to aquarium-cultured Aurelia sp. from Japan. They differ consistently in radial canal morphology. In allozyme electrophoresis, significant differences at 12 of 14 polymorphic loci strongly suggest that Aurelia sp. from Foster City and Tokyo Bay belong to one species, while Aurelia sp. from Monterey Bay and Vancouver Island belong to a second species. We propose that Aurelia sp. at Foster City is a recent introduction, possibly from Japan via ships' ballast water. The identities and taxonomic affinities of the two Aurelia defined in this study, and their relationships with the Linnaen A. aurita described from the North Atlantic, will require genetic and morphological study of the currently recognized species A. aurita and A. limbata (Brandt, 1838) from several zoogeographical provinces.     

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.     

Density-dependent effects control the reproductive strategy and population growth of <Emphasis Type="Italic">Aurelia</Emphasis><Emphasis Type="Italic">aurita</Emphasis> s.l. scyphistomae

Aureliaaurita s.l. scyphistomae are capable of developing different asexual modes for propagation and thus present a multi-mode reproductive strategy. The reproduction rates and the reproductive strategy they adopt depend on a combination of various environmental parameters. We investigated the A.aurita s.l. polyp-to-polyp reproduction strategy and population growth in relation to polyp density. Our results confirmed that density-dependent factors control population growth of A. aurita s.l. scyphistomae in three different ways: (1) decreasing the polyp reproduction rate, (2) triggering the production of motile bud-like tissue particles and (3) inducing the detachment of developed scyphistomae. Whereas the decrease in the reproduction rate reduces the number of recruits, the motile particles and the detachment of scyphistomae contribute to minimizing density-dependent effects by allowing reproductive products and scyphistomae to drift away. Thus, not only are the negative effects of intraspecific competition for space and food diminished but also the potential colonization of new substrates, and further increase in scyphistoma density is favoured on larger spatial scales. The potential capability to switch its polyp-to-polyp reproduction strategy in response to environmental clues and population density may give Aurelia high adaptability in the temperate coastal waters where they commonly live and where they may be exposed to wide-ranging and fluctuating environmental variables that affect their survival and longevity. Considering these features, as well as the wide tolerance of Aurelia scyphistomae (and medusae) to environmental parameters, it is not surprising that the species/lineages of Aurelia are cosmopolitan and exhibit the most frequent bloom events worldwide.     

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.     

Genetic analysis of sexual reproduction in the dendrophylliid coral Balanophyllia elegans

The ahermatypic scleractinian Balanophyllia elegans has served as a model of limited larval dispersal in an aclonal species. However, other species from the same family (Dendrophylliidae) produce larvae asexually, and closely connected polyps of B. elegans, potentially the result of asexual reproduction, are commonly observed in the field. Here, we use a combination of laboratory experiments, controlled crosses, and genetic surveys of field-collected individuals to demonstrate (1) marker allozymes are inherited in accordance with Mendel's laws, (2) polyps that are connected in the field are not genetically identical, and (3) laboratory manipulation of a single polyp can produce a second polyp on the aboral side of the original, but not adjacent to the original. In combination, these results suggest that the larvae of B. elegans result from mictic sexual reproduction, and that connected polyps result not from asexual budding but rather from the fusion of genetically distinct individuals.     

Comparison of telomere length among different life cycle stages of the jellyfish Cassiopea andromeda

The polyp (scyphistoma) of the jellyfish Cassiopea andromeda reproduces asexually repeatedly, while the medusa, the sexually reproducing stage, exhibits a relatively shorter life span. As a first step to understand the mechanism behind the differences in the life spans of the polyp and medusa stages of the jellyfish, we compared the lengths of the telomere region of one targeted chromosome between the polyp and medusa stages using a modified single telomere length assay (STELA). The double-stranded regions of the telomeres were amplified by PCR, and the average length of the PCR products was estimated by densitometry analysis of the gel smear. Chromosomes within cells of the bell region of the medusa were characterized by longer telomeres than those of polyps, asexual propagules, or other regions of the medusa. This is the first study to estimate the telomere lengths of targeted chromosomes in a cnidarian and opens a way to understand the mechanism underlying different life spans of the polyp and medusa stages.     

Determination of δ13C and δ15N and trophic fractionation in jellyfish: implications for food web ecology

Application of stable isotope analysis (SIA) in jellyfish allows definition of trophic patterns not detectable using gut content analysis alone, but analytical protocols require standardization to avoid bias in interpreting isotopic data. We determined δ¹³C and δ¹⁵N in Aurelia sp. from the northern Gulf of Mexico (30°00′N, 89°00′W–30°24′N, 88°00′W) to define differences in stable isotope composition between body parts and whole body, the effect of lipid extraction on δ¹³C in tissues, and fractionation values from medusa to prey. The isotopic composition of bell and whole Aurelia sp. was not different. The increase in δ¹³C values after lipid removal suggested a correction is needed. To aid future analyses, we derived a correction equation from empirical data for jellyfish samples. Laboratory feeding experiments indicated medusae increased +4 ‰ in δ¹³C and +0.1 ‰ in δ¹⁵N compared to their diet. These results suggest protocols commonly applied for other species may be inaccurate to define Aurelia sp. trophic ecology. Because Aurelia spp. are commonly found in marine ecosystems, accurately defining their trophic role by use of SIA has implications for understanding marine food webs worldwide.     

Asexual repooduction in Glossobalanus crozieri (Ptychoderidae,Enteropneusta, Hemichordata)

Several aspects of asexual reproduction in Glossobalanus crozieri (a species first found in Bermuda, but occurring also on the coast of Brazil) have been observed. The results are compared with those by earlier authors on other species of Enteropneusta. A revision of asexual reproduction in the Enteropneusta (Hemichordata) is made. Developmental rates and regeneration were studied in branchiogenital individuals, hepatic individuals and regenerands. The habitat where these forms are found is described. Possible meaning of gonad maturation and sexual reproduction in relation to asexual reproduction in different species is discussed. The need for further studies on the variation of forms which reproduce both sexually and asexually is emphasized, with a view to taxonomic problems, based both upon the proportions of the different body regions and on the number of branchial pores. The present findings indicate that, when compared to the results reported for other species, asexual reproduction in all ptychoderids (Balanoglossus, Glossobalanus and Ptychodera) follows the same general pattern. The phenomenon of asexual reproduction in this group is also confirmed to be worldwide.     

Evidence for broadcast spawning as well as brooding in the scleractinian coral Pocillopora damicornis

The reproductive activity of 88 colonies of Pocillopora damicornis at Rottnest Island, Western Australia, was examined over 12 mo (20 August 1988 to 20 August 1989). Larvae were found close to the three new moons of February, March and April, 1989, but in large numbers only during March; 67% of all corals produced larvae. There was no obvious connection between gametogenesis and the production of brooded larvae; however, those corals which did not produce, larvae also did not produce ova but did produce, sperm, and must be considered male colonies. All colonies which produced ova went on to produce planulae. Spermatogenesis occurred throughout the year, but oogenesis occurred between September 1988 and April 1989. Dramatic declines in the percentage of polyps containing eggs and sperm occurred in early February and early April 1989. On these dates the amount of lipid in the tissue of the corals decreased, which suggests that the decline in the percentage of polyps with sperm and eggs was not due to the gametes being resorbed by the polyps. Therefore, P. damicornis at Rottnest Island appears to be both a brooder and a broadcaster with asexual reproduction through brooded larvae and sexual reproduction through spawning of gametes. This accounts for earlier evidence in the literature that some input from sexual reproduction had been occurring in the populations of P. damicornis at Rottnest Island. There is no direct evidence for spawning, as broadcasting of gametes has not been observed, but the indirect evidence provides a strong argument for the existence of this dual reproduction.     

Complex reproductive cycle inEucheilota paradoxica (Hydrozoa: Leptomedusae): Medusae,polyps and frustules produced from medusa stage

Eucheilota paradoxica Mayer, 1900 is a small leptomedusa unknown in the Mediterranean Sea until 1977. Since then, it has become endemic and occurs in great abundance in autumn in the bay of Villefranche-sur-mer. Since no sexual stages have ever been collected, either from the Atlantic Ocean or from the Mediterranean Sea, we decided to investigate the still unknown life cycle of this species. Specimens were obtained in the bay of Villefranche-sur-mer from plankton hauls between 0 and 50 m depth during October and November 1986. At temperatures >18°C, the medusa reproduces intensively by asexual medusal budding (one young medusa liberated per two days). This asexual reproduction explains the rapid proliferation of the species and the origin of the large population every autumn. In cultures maintained at temperatures <18°C, two other reproduction modes, not yet reported in a leptomedusa, were observed: (1) At 16° to 17°C, frustules bud along the canals; these frustules are dormant, giving rise to a small polyp when the temperature increases once more above 18°C. (2) At temperatures close to 15°C each medusa, in place of the gonads, differentiate into one or two polyps; these polyps develop progressively, concomitant with gradual regression of the mother medusa.     

Direct and indirect trophic interactions of <Emphasis Type="Italic">Aurelia </Emphasis>sp. (Scyphozoa) in a stratified marine environment (Mljet Lakes,Adriatic Sea)

The pattern of diel vertical migration and the trophic interactions of moon jelly (Aurelia sp.) were investigated in the sea lakes of Mljet Island (Adriatic Sea) where this scyphomedusa is present throughout the year. Water column characteristics, plankton and in situ behaviour of Aurelia were followed over several 24-h cycles (6–8 times during each cycle) from the surface to the bottom (44 m). Aurelia exhibited a consistent pattern of diel vertical migration. Most of the time Aurelia were located at the bottom of the thermocline layer at temperatures lower than 19°C. Aurelia migrated towards the surface at dusk when the majority was found within the thermocline or just above it. During the night the medusae sank into the deepest layers below 25 m. The main medusa food items inferred from stomach contents were small adult copepods like Oithona nana and Paracalanus parvus and copepodites of small calanoids and cyclopids. In addition, in situ feeding experiments indicated high clearance rates for nauplii and naked ciliates and clear response of bacterial populations pointing to indirect cascade effects of Aurelia on microbial in addition to classical food web.     

Allozyme electrophoresis demonstrates that the scleractinian coral Montipora digitata is two species

Allozyme electrophoresis at five loci was used to clarify the species status of two morphs of the scleractinian coral Montipora digitata (Dana, 1846), using specimens collected from three locations off the Queensland coast between October 1991 and April 1993. The two morphs, occurring sympatrically, were distinguished by one fixed gene difference at Locus LT-2 and frequency differences at three other loci, indicating reproductive isolation between them. Populations of both morphs were generally in Hardy-Weinberg equilibrium, suggesting that sexual reproduction does occur. Despite sampling designed to reduce the chance of collecting clonemates, the genetic diversity ratio (G _O:G_E) indicated that asexual reproduction also occurs. This is in agreement with the known modes of reproduction for this species. There was significant genetic differentiation between populations of one of the morphs of M. digitata. This could have been influenced by greater degrees of asexual reproduction in this morph, selection within sites, or variation in larval survival and recruitment patterns. Relatively low F _st (differentiation among populations) values found for a broadcast spawning species during this study are in accordance with the hypothesis that brooded planulae are adapted for rapid settlement, whereas planulae produced by broadcast spawners are adapted for widespread dispersal.     

Effect of temperature,salinity and food level on sexual and asexual reproduction in Brachionus plicatilis (Rotifera)

The reproductive response of sexual and asexual female Brachionus plicatilis (Muller) was examined over temperatures ranging from 20° to 40°C, salinities from 5 to 40 S, and food levels from 0.25 to 20 g Chlorella vulgaris dry-weight per ml. Reduced food levels, as well as temperature and salinity extremes, reduced reproduction of both sexual and asexual females, but did so differentially. Reproduction by sexual females was reduced to a greater extent at environmental extremes than asexual females. The broad, flat reproductive response curve of asexual females extended beyond the limits of the narrower, more sharply peaked curve of sexual females. Thus zones of exclusively asexual reproduction exist at environmental extremes where sexual reproduction is physiologically restricted. These results are corroborated by a comparison of the lifetime fecundity of individual sexual and asexual females over a 20°C temperature range. No differences in lifetime fecundity occurred between sexual and asexual females at 18° and 28°C. At 38°C, however, asexual female fecundity reached its highest level, while sexual female fecundity declined 15%. The appearance of sexual females in rotifer populations in the result of both inducible and repressible factors.     

Effects of temperature,salinity, and predators on mortality of and colonization by the invasive hydrozoan<Emphasis Type="Italic"> Moerisia lyonsi</Emphasis>

The successful invasion of non-indigeneous species depends on initial colonization as well as establishing a self-maintaining population. The invasive hydrozoan Moerisia lyonsi (Boulenger, 1908), possibly originating from low-salinity waters in the Black Sea and Middle East regions, has become established in low-salinity waters in several estuaries of North America, including Chesapeake Bay. The effects of temperature and salinity on mortality of M. lyonsi polyps were examined in the laboratory in February 2001 in the presence of abundant food. The polyps of M. lyonsi were directly transferred from 20°C and 10 salinity to one of 45 combinations of temperature (10–29°C) and salinity (1–40). Polyp mortality within 7 days occurred only in low-temperature treatments with salinities of 35–40. Surviving polyps reproduced asexually in salinities of 1–40 at 20–29°C, and in salinities of 1–25 at 15°C, but not in any salinities at 10°C. The greatest asexual reproduction rates, an index for population survival potential, occurred at salinities of 5–20. Survival and reproduction of M. lyonsi over such broad temperature and salinity ranges indicate that M. lyonsi may colonize and establish populations throughout the Chesapeake Bay; however, M. lyonsi medusae were reported only at salinities <9.3 there. This discrepancy may be due to the effects of predators. The scyphomedusan Chrysaora quinquecirrha (Desor, 1848), but not the ctenophore Mnemiopsis leidyi (A. Agassiz, 1865) consumed M. lyonsi medusae in laboratory experiments in August–September 2001. Populations of M. lyonsi do not appear to be limited by temperature and salinity conditions; however, their distribution in Chesapeake Bay may be restricted to low salinities not inhabited by predators.Communicated by J.P. Grassle, New Brunswick     

Reproductive strategies of Mycalecontarenii (Porifera: Demospongiae)

The sexual and asexual phases of the reproductive cycle of the sponge Mycalecontarenii (Martens, 1824) were studied from samples collected over a period of 2 years (June 1994 to May 1996) in a Mediterranean coastal basin (Porto Cesareo, southwestern Apulia). The species is viviparous; year round it shows a discontinuous oocyte production. No males were found, possibly owing to the very short period of spermatogenesis. M.␣contarenii produces asexual buds during autumn and winter, and this process seems to be related to changes in water temperature. Even though, at present, we cannot evaluate the incidence of larval recruitment, we postulate that asexual reproduction in M.contarenii plays a crucial role in maintaining the sponge population. Received: 8 July 1997 / Accepted: 5 November 1997     

Differences between native and invasive<Emphasis Type="Italic"> Caulerpa taxifolia</Emphasis>: a link between asexual fragmentation and abundance in invasive populations

The tropical and subtropical marine green alga Caulerpa taxifolia has invaded several temperate regions throughout the world, including southeastern Australia. In this study, I examined how invasive C. taxifolia from temperate southeastern Australia differed from native C. taxifolia from subtropical Moreton Bay, Australia, in the traits proposed as being important to its invasion success: thallus size and density, levels of asexual reproduction (fragmentation) and total biomass. Against the prediction of a large size for invasive C. taxifolia, native populations from Moreton Bay had larger stolons and fronds than invasive populations. However, invasive populations consistently had much higher densities of stolons, fronds and fragmented fronds; and a greater biomass compared to native populations. Average densities at invasive sites exceeded 4,700 stolons and 9,000 fronds/m² and were as high as 27,000 stolons and 95,000 fronds/m², which are the highest reported for C. taxifolia anywhere. Average densities of fragmented fronds at invasive sites were as high as 6,000/m² and up to 45% of all stolons at invasive sites could be directly linked to asexual recruitment via fragmented fronds. Importantly, at invasive locations there was a strong association between asexual reproduction and abundance demonstrated by positive correlations between the density of fragmented fronds and total biomass. These findings are the first to describe quantitative differences between native and invasive C. taxifolia and to demonstrate a link between the high levels of asexual reproduction and high abundance in invasive populations. Although the causes and consequences of high levels of asexual reproduction remain to be explored, this study suggests that changes in demographic and life-history traits during the invasion by C. taxifolia into temperate habitats may contribute to its success there.Communicated by M. S. Johnson, Crawley     

Population genetic structures of the fissiparous seastar Coscinasterias acutispina in the Sea of Japan

The morphological characteristics and the population genetic structures of the fissiparous seastar Coscinasterias acutispina were investigated for eight sites in the Sea of Japan in order to clarify the presence of sexual and asexual reproduction. Morphological observation based on arm length showed that fission was common at all eight sites examined, indicating the likely production of clonal individuals. A random amplified polymorphic DNA (RAPD) marker was used to detect clones arising by fission and to assess gene flow among sites. A simulation approach using RAPD data revealed the presence of clonal individuals at almost all sites, suggesting the existence of asexual reproduction. The result of phylogenetic analysis according to RAPD genotype showed no relationship between genetic and geographic distances. Considering the limited movement ability of seastar species during the adult phase, these observations suggest the existence of marked gene flow among sites, due to dispersal of planktonic larvae produced by sexual reproduction. These observations suggest that multi-locus genotypic compositions depend on the relative amounts of recruitment from sexual and asexual reproduction in each population.     

Genetic variability and differentiation of sporophytes and gametophytes in populations of Gelidium arbuscula (Gelidiaceae: Rhodophyta) determined by isozyme electrophoresis

Genetic differentiation and genetic variability of sporophytic and gametophytic populations of Gelidium arbuscula (Bory) from three localities sampled in 1989 and 1990 in the Canary Islands (Spain) were examined by isozyme electrophoresis. Twenty-three to 29 putative alleles corresponding to 22 gene loci, were compared. High deviations in Hardy-Weinberg equilibrium, and significant differences between allelic frequencies of sporophytic and gametophytic subpopulations at the same locality were found, suggesting a predominant asexual reproduction of G. arbuscula. The genetic variability (percentage of polymorphic loci, mean number of alleles per locus and average gene diversity) of haploid subpopulations was lower than that of diploid subpopulations at all three localities, being the lowest described for seaweeds. No correlation between genetic and geographical distance was found. The high genetic differentiation coefficient between all subpopulations suggests a very reduced genetic flow between subpopulations of the same and of different localities. These results suggest that the genetic structure of the populations of G. arbuscula from the Canary Islands is due to a founder-effect combined with a predominance of asexual reproduction. This is the first report comparing allelic frequencies between sporophytic and gametophytic subpopulations of seaweeds.