Digenea parasites of jellyfish and ctenophores of the southern Atlantic

Parasites of planktonic cnidarians and ctenophores in the southern Atlantic Ocean are little known. The aim of this study was to describe three new metacercariae from jellyfish and ctenophores, and assess the importance of the gelatinous zooplankton as intermediate hosts in the life history of digeneans. During examination of zooplankton in Argentine Sea for digeneans that mature in fishes, two species of jellyfish (Phialidium sp. and Liriope tetraphylla Chamiso & Eysenhardt, 1821), and one ctenophore (Mnemiopsis mccradyi Mayer, 1900) were analyzed for parasites. The samples were obtained in Mar del Plata. Three metacercariae belonging to Faustulidae, Lepocreadiidae and Hemiuridae are described. The prevalence (percent of hosts infected) varied from 1.4–30% and the range of intensity (number of individuals of a parasite species in a single infected host) was from 1 to 30 for the different metacercariae. Given the important position of free-swimming cnidarians and ctenophores in the marine food web, and the great number of fishes that have been found with these organisms in their digestive tracts, their importance in the life histories of digeneans should not be underrated.     

Gelatinous zooplankton of the Arctic Ocean: in situ observations under the ice

The gelatinous zooplankton of the Canada Basin were observed with a deep-diving remotely operated vehicle (ROV) during August–September, 2002. Taxa observed fell into four main groups: cnidarians, ctenophores, chaetognaths, and pelagic tunicates. We provide detailed data on the vertical distributions of many taxa from three sites which span the Canada Basin. The most common gelatinous organisms in the surface waters were the ctenophores Mertensia ovum and Bolinopsis infundibulum. These two species were found in very large numbers in the near-surface mixed layer. In the mesopelagic zone, below the transition from the Pacific water layer and the Atlantic water layer, the most common species was Sminthea arctica. Surprising numbers of the scyphomedusa Atolla tenella were found in the deep waters of the basin, along with an undescribed species of narcomedusae. The vertical distributions of the gelatinous zooplankton observed with the ROV show several trends related to the physical properties of the water and geographic location within the basin.     

Seasonal variation of gelatinous plankton consumption by fish in the South-western Atlantic Ocean: a question of strategy*

Gelatinous plankton is an important food resource for several species of fishes in the Southwestern Atlantic Ocean. Some fish depend heavily on these organisms and are specialized to feed on ctenophores, salps and medusae, while others only consume gelatinous plankton occasionally. We hypothesize that consumption of gelatinous plankton by fishes represents an alternative food resource when primary prey are not available during cold periods in the study area. To determine seasonal variations of gelatinous plankton consumption by fishes, data samples were grouped into cold and warm periods. A total of 64,567 stomachs belonging to 106 species of fish were analysed, of which 32,943 (51%) contained food items. Of those containing food items, 2719 (8.2%), from 38 fish species, contained gelatinous items. Fourteen species ingested gelatinous in warm period, nine in cold period, and 15 in both periods. The proportion of stomachs with gelatinous was significantly greater during the cold period. Ctenophores were the most predominant prey in both periods, followed by salps and medusae. Consumption of ctenophores, salps and medusae was unevenly distributed within the area during the different periods. Classification methods (group average sorting utilizing Bray–Curtis similarity measures based on log (X + 1) identified eight areas of consumption. SIMPER (similarity percentages) analyses revealed that nine fish species contributed most to gelatinous plankton consumption. The seasonal and spatial variation of gelatinous consumption by fish would be related to the availability of food in each period. Strategies of gelatinous consumption, including survival, feeding opportunities and prey specialization, are discussed.     

A comparison of carbon-specific respiration rates in gelatinous and non-gelatinous zooplankton: A search for general rules in zooplankton metabolism

Using 470 data from the literature the dry weight-specific respiration rates of gelatinous zooplankton (cnidarians, ctenophores and salps) and non-gelatinous zooplankton (mainly crustacea) were converted to carbon-specific values. The resulting carbon-specific respiration rates showed no significant differences between the two groups of zooplankton, indicating similar oxygen requirements per gram of carbon biomass. From this finding, it can be suggested that the differences in the rates of oxygen consumption measured in the two types of zooplankton in the sea can be explained by the carbon biomass ratio between gelatinous and non-gelatinous zooplankton. Furthermore, the low rate of metabolism of gelatinous species compared with that of non-gelatinous animals of the same volume can be attributed predominantly to the relatively low organic matter content in the former. It is recommended that all weight-specific metabolism rates be expressed using carbon as body mass unit (e.g. mg O₂ gC⁻¹ d⁻¹) which enables more accurate comparisons between individuals exhibiting different dry weight/carbon ratios.     

New records of Amphipoda Hyperiidea in associations with gelatinous zooplankton

Associations between Amphipoda Hyperiidea specimens and gelatinous zooplankton such as cnidarians, ctenophores or salps are known in the literature. It is well established that many hyperiids are intimately associated with gelatinous zooplankton during part or all of their life-cycles, and that the associations could be of commensal or of parasitic nature, sometimes very highly species-specific. Although conventional net sampling usually does not preserve evidence of these associations, 12 instances of association (I.A.) were found in nine of the 49 samples collected during the 'FINEP XI Cruise', carried out on the Southeast coast of Brazil (23–28 °S), in January 1980. Lestrigonus bengalensisGiles, 1887 (11 specimens) and Simorhynchotus antennarius(Claus, 1871) (one specimen) were found inside salps and siphonophores. The study of the nature of these associations is essential to the understanding of the evolution of the group.     

The Development of Radial and Biradial Symmetry: The Evolution of Bilaterality

SYNOPSIS. Understanding the evolutionary origin of novel metazoanbody plans continues to be one of the most sought after answersin biology. Perhaps the most profound change that may have occurredin the Metazoa is the appearance of bilaterally symmetricalforms from a presumably radially symmetrical ancestor. The symmetryproperties of bilaterally symmetrical larval and adult metazoansare generally set up during the cleavage period while most "radially"symmetrical cnidarians do not display a stereotyped cleavageprogram. Ctenophores display biradial symmetry and may representone intermediate form in the transition to bilateral symmetry.The early development of cnidarians and ctenophores is comparedwith respect to the timing and mechanisms of axial determination.The origin of the dorsal-ventral axis, and indeed the relationshipsof the major longitudinal axes, in cnidarians, ctenophores,and bilaterian animals are far from certain. The realizationthat many of the molecular mechanisms of axial determinationare conserved throughout the Bilateria allows one to formulatea set of predictions as to their possible role in the originsof bilaterian ancestors.     

From gelatinous to muscle food chain: rock cod Patagonotothen ramsayi recycles coelenterate and tunicate resources on the Patagonian Shelf

Stomach contents of 4808 fishes of 20 species caught in the eastern part of the Patagonian Shelf between 1999 and 2012 were analysed to assess dietary contributions of gelatinous plankton resources. Gelatinous plankton occurred in diets of seven species with two species, Patagonotothen ramsayi and Squalus acanthias, having >10% ctenophores in their diet. Consumption of gelatinous plankton was important in P. ramsayi and was strikingly seasonal, with maximum occurrence (up to 46% of non‐empty stomachs) in late summer to autumn. Ctenophores were most abundant in P. ramsayi of 25–34 cm total length, L_T, whereas salps were more frequent in larger >35 cm L_T individuals. In winter to spring, occurrence of gelatinous plankton in diets was minimal, reflecting their overall seasonal abundance in the ocean. The recent increase in abundance of P. ramsayi has enabled the species to recycle a significant proportion of the ecosystem production from gelatinous dead end to the main muscular food chain via seasonal reliance on ctenophores, jellyfish and tunicates. This additional influx of production that has been diverted from the gelatinous food chain favours the increase in abundance of several piscivorous top predators and affects the trophic web structure of the Patagonian Shelf ecosystem.     

Interactions of pelagic cnidarians and ctenophores with fish: a review

Medusae, siphonophores and ctenophores (here grouped as `pelagic coelenterates') interact with fish in several ways. Some interactions are detrimental to fish populations, such as predation by gelatinous species on pelagic eggs and larvae of fish, the potential competition for prey among pelagic coelenterates and fish larvae and zooplanktivorous fish species, and pelagic coelenterates serving as intermediate hosts for fish parasites. Other interactions are positive for fish, such as predation by fish on gelatinous species and commensal associations among fish and pelagic coelenterates. The interactions range from beneficial for the gelatinous species (food, parasite removal), to negative (predation on them). We review existing information and present new data on these topics. Although such interactions have been documented frequently, the significance to either fish or pelagic coelenterate populations is poorly understood. The effects of pelagic coelenterates on fish populations are of particular interest because of the great importance of fisheries to the global economy. As fishing pressures mount, it becomes increasingly important to understand how they may influence the balance between pelagic coelenterates and fish.     

Associations between gelatinous zooplankton and hyperiid amphipods (Crustacea: Peracarida) in the Gulf of California

Hydrobiologia - Hyperiid amphipods are pelagic crustaceans that live associated with gelatinous zooplankton including medusae, ctenophores, siphonophores, and salps. Standard plankton sampling...     

Gelatinous zooplankton (ctenophores,salps and medusae): an important food resource of fishes in the temperate SW Atlantic Ocean

This study quantifies the occurrence of gelatinous zooplankton in the stomach contents of fishes from the southwest Atlantic Ocean (33°–55°S). More than 69,000 fish stomachs belonging to 107 species were examined. A total of 39 fishes were documented as consumers of gelatinous zooplankton, 23 of which were newly discovered. Three gelatinous organism consumption categories are recognized: (1) very frequent consumers (10 species, six of which were exclusive); frequent consumers (five species); and occasional consumers (26 species). Three types of gelatinous prey (ctenophores, salps and medusae) were found in the stomach contents of fishes. Ctenophores were consumed at high levels across almost the entire continental shelves of Argentina and Uruguay. Salps were frequent prey on the slope and southern shelf. In contrast, medusae were consumed in coastal areas, slopes and the southern shelf. Classification methods (group average sorting of the Bray–Curtis similarity measures based on log (X?+?1)-transformed percentage data) determined six areas where fishes predated on gelatinous organisms. SIMPER (similarity percentages) analysis determined which fishes contributed more to the consumption of gelatinous organisms. Results revealed that two fish species (Stromateus brasiliensis and Squalus acanthias) had high gelatinous zooplankton predation rates throughout the entire study area, while another six species (Patagonotothen ramsayi, Helicolenus dactylopterus lahillei, Macrourus holotrachys, Merluccius hubbsi, Schroederichthys bivius, and Macruronus magellanicus), while widely distributed, seemed to have specific areas where consumption occurred. This study not only provides new knowledge about the importance of gelatinous zooplankton in the diet of numerous fishes, but might also be valuable for planning and managing local fisheries.     

A molecular phylogenetic framework for the phylum Ctenophora using 18S rRNA genes.

This paper presents the first molecular phylogenetic analysis of the phylum Ctenophora, by use of 18S ribosomal RNA sequences from most of the major taxa. The ctenophores form a distinct monophyletic group that, based on this gene phylogeny, is most closely related to the cnidarians. Our results suggest that the ancestral ctenophore was tentaculate and cydippid-like and that the presently recognized order Cydippida forms a polyphyletic group. The other ctenophore orders that we studied (Lobata, Beroida, and Platyctenida) are secondarily derived from cydippid-like ancestors, a conclusion that is also supported by developmental and morphological data. The very short evolutionary distances between characterized ctenophore 18S rRNA gene sequences suggests that extant ctenophores are derived from a recent common ancestor. This has important consequences for future studies and for an understanding of the evolution of the metazoans.     

Myxosporea (Myxozoa,Cnidaria) Lack DNA Cytosine Methylation

DNA cytosine methylation is central to many biological processes, including regulation of gene expression, cellular differentiation, and development. This DNA modification is conserved across animals, having been found in representatives of sponges, ctenophores, cnidarians, and bilaterians, and with very few known instances of secondary loss in animals. Myxozoans are a group of microscopic, obligate endoparasitic cnidarians that have lost many genes over the course of their evolution from free-living ancestors. Here, we investigated the evolution of the key enzymes involved in DNA cytosine methylation in 29 cnidarians and found that these enzymes were lost in an ancestor of Myxosporea (the most speciose class of Myxozoa). Additionally, using whole-genome bisulfite sequencing, we confirmed that the genomes of two distant species of myxosporeans, Ceratonova shasta and Henneguya salminicola, completely lack DNA cytosine methylation. Our results add a notable and novel taxonomic group, the Myxosporea, to the very short list of animal taxa lacking DNA cytosine methylation, further illuminating the complex evolutionary history of this epigenetic regulatory mechanism.     

Expansion of the SOX gene family predated the emergence of the Bilateria

Members of the SOX gene family are involved in regulating many developmental processes including neuronal determination and differentiation, and in carcinogenesis. So far they have only been identified in species from the Bilateria (deuterostomes and protostomes). To understand the origins of the SOX family, we used a PCR-based strategy to obtain 28 new sequences of SOX gene HMG domains from four non-bilaterian Metazoa: two sponge species, one ctenophore and one cnidarian. One additional SOX sequence was retrieved from EST sequences of the cnidarian species Clytia hemisphaerica. Unexpected SOX gene diversity was found in these species, especially in the cnidarian and the ctenophore. The topology of gene relationships deduced by Maximum Likelihood analysis, although not supported by bootstrap values, suggested that the SOX family started to diversify in the metazoan stem branch prior to the divergence of demosponges, and that further diversification occurred in the eumetazoan branch, as well as later in calcisponges, ctenophores, cnidarians and vertebrates. In contrast, gene loss appears to have occurred in the nematode and probably in other protostome lineages, explaining their lower number of SOX genes.     

Cloning,Sequencing, Expression and Structural Investigation of Mnemiopsin from <Emphasis Type="Italic">Mnemiopsis leidyi</Emphasis>: An Attempt Toward Understanding Ca<Superscript>2+</Superscript>-Regulated Photoproteins

A comparison of the two most famous groups of calcium-regulated photoproteins, cnidarians and ctenophores, showed unexpectedly high degree of structural similarity regardless of their low sequence identity. It was suggested these photoproteins can play an important role in understanding the structural basis of bioluminescence activity. Based on this postulate, in this study the cDNA of mnemiopsin from luminous ctenophore Mnemiopsis leidyi was cloned, expressed, purified and sequenced. The purified cDNA, with 621 base pairs, coded a 206 residues protein. Sequence of mnemiopsin showed 93.5 and 51% similarity to other ctenophore proteins and cnidarians, respectively. The cDNA encoding apo-mnemiopsin of M. leidyi was expressed in Escherichia coli. The purified apo-protein showed a single band on SDS-PAGE (molecular weight ~27 kDa). A semi-synthetic mnemiopsin was prepared using coelenterazine and EDTA and its luminescence activity was measured in the presence of CaCl₂. The results showed an optimum pH of 9.0 and lower calcium sensitivity compared to aequorin. Comparison of amino acid residues in substrate binding site indicated that binding pocket of ctenophores contains less aromatic residues than cnidarians. This can lead to a decline in the number of stacking interactions between substrate and protein which can affect the stability of coelenterazine in binding cavity. Structural comparison of photoproteins with low sequence identity and high 3D structural similarity, can present a new insight into the mechanism of light emission in photoproteins.     

A golden age of gelata: past and future research on planktonic ctenophores and cnidarians

The study of the natural history of gelatinous zooplankton (‘gelata’) reached a high point at the end of the 19th century, when scientists first began to understand the phylogenetic and ecological links between cnidarians and ctenophores. Siphonophores, carefully figured in their entirety, and gauze-like lobate ctenophores too fragile to touch, were described by the dozens. In the ensuing years, focus on zooplankton shifted toward more ‘industrial’ goals such as quantitative sampling using plankton nets. While plankton scientists were busy summing tattered parts, they lost sight of the whole jellies themselves, and a crustaceocentric view of the ocean came to dominate. During this period, the most dramatic breakthroughs in cnidarian research came from laboratory studies of neurobiology, physiology, and development, particularly of certain model organisms. Now, at the turn of this century, we have the opportunity to bring gelata back into primacy. Submersibles and remotely operated vehicles allow us to study entire life histories of organisms that we did not even know existed. The tools of molecular biology allow us to answer questions about development, evolution, and phylogeny that had reached a stalemate. Even in the surface waters, where it might be thought that there is little left to learn, in situ observations have revealed unexpected interactions and hidden diversity. The critical roles that these organisms play in the health of the oceans, their position at the crux of many evolutionary debates, and the tools for biotechnology that they provide, have led to resurgent public appreciation and awareness. Although advanced tools do not necessitate good science, we have few excuses for failing to bring about another golden age of gelata. A plenary address of the 7th International Conference on Coelenterate Biology.     

Associations between gelatinous zooplankton and hyperiid amphipods (Crustacea: Peracarida) in the Gulf of California

Hyperiid amphipods are pelagic crustaceans that live associated with gelatinous zooplankton including medusae, ctenophores, siphonophores, and salps. Standard plankton sampling disrupts natural associations, so the most reliable way to determine an association is through direct observation of the organisms in their environment. The planktonic fauna of the Gulf of California dwelling between 10 and 3000 m was surveyed using SCUBA diving and a remotely operated submersible (ROV) during March 2003. Here we report our observations on a total of 14 symbiotic associations found between the hyperiid amphipods and various taxa of gelatinous zooplankton. We found parental care behavior in a group of amphipods (Oxycephalidae) in which this phenomenon has not been previously reported. For two hyperiid species, Euthamneus rostratus and Vibilia australis, we present the first information on their symbiotic relations. Additional hosts were discovered for other well-known and widely distributed hyperiid species (i.e. Brachyscelus crusculum, Hyperoche medusarum). Photographic evidence of some of these interactions is included in this contribution. This is the first survey of these relationships in the Gulf of California, and many aspects of the ecology and biology of these symbioses remain to be studied.     

Haeckelia (= Euchlora) and Hydroctena and the phylogenetic interrelationships of Cnidaria and Ctenophora

A scrutiny of the literature shows that the ctenophore Haeckelia (= Euchlora) ruba has only kleptocnidae and that Hydroctena salenskii is a ctenophore without special cnidarian affinities. The “missing links” between cnidarians and ctenophores have thus turned out to be based on misinterpretations and must be excluded from future discussions on phylogeny.     

Trophic ecology of planktonic gelatinous predators in Saanich Inlet, British Columbia: diets and prey selection

The diets of most of the gelatinous predators (medusae and ctenophores)in Saanich Inlet, British Columbia (0–25 m) were dominatedduring the spring by eggs of Euphausia pacifica which formed     

Does the nutrient stoichiometry of primary producers affect the secondary consumer Pleurobrachia pileus?

We investigated whether phosphorus limitations of primary producers propagate upwards through the food web, not only to the primary consumer level but also onto the secondary consumers’ level. A tri-trophic food chain was used to assess the effects of phosphorus-limited phytoplankton (the cryptophyte Rhodomonas salina) on herbivorous zooplankters (the copepod Acartia tonsa) and finally on zooplanktivores (the ctenophore Pleurobrachia pileus). The algae were cultured in phosphorus-replete and phosphorus-limited media before being fed to two groups of copepods. The copepods in turn were fed to the top predator, P. pileus, in a mixture resulting in a phosphorus-gradient, ranging from copepods having received only phosphorus-replete algae to copepods reared solely on phosphorus-limited algae. The C:P ratio of the algae varied significantly between the two treatments, resulting in higher C:P ratios for those copepods feeding on phosphorus-limited algae, albeit with a significance of 0.07. The differences in the feeding environment of the copepods could be followed to Pleurobrachia pileus. Contrary to our expectations, we found that phosphorus-limited copepods represented a higher quality food source for P. pileus, as shown by the better condition (expressed as nucleic acid content) of the ctenophore. This could possibly be explained by the rather high C:P ratios of ctenophores, their resulting low phosphorus demand and relative insensitivity to P deficiency. This might potentially be an additional explanation for the observed increasing abundances of gelatinous zooplankton in our increasingly phosphorus-limited coastal seas.