The influences of ecto- and endoderm in determining the axial polarity ofHydra attenuata Pall. (Cnidaria,Hydrozoa)

Summary Ectoderm and endoderm of the gastric column ofHydra attenuata Pall. were separated from each other and reassembled with either the same (controls) or opposite polar orientation. The controls always regenerated a head and basal disc according to the original polarity. In those specimens in which the polarity of ecta- and endoderm was opposite 33 specimens out of 41 reconstituted a single polyp whose body axis was clearly identifiable. Of these cases 8 followed the polarity of the ectoderm, 3 obeyed that of the endoderm, and 22 formed axes perpendicular to the original longitudinal axis. In 5 cases 2 specimens regenerated from the reassembled specimens. It is thus demonstrated that axial polarity ofHydra is determined by both the ecto- and the endoderm.This work was carried out with the support of the Swiss National Science Foundation (Grant Nr. 3.317-0.78)     

Structure and function of the locomotory system ofPolyorchis montereyensis (Cnidaria,Hydrozoa)

Summary 1. The structure and function of the locomotory system of the anthomedusanPolyorchis montereyensisSkogsberg were studied in detail. Anatomical investigations were carried out primarily on fresh or formalin fixed specimens; histology was done on specimens fixed in Bouin's fluid. Functional analyses were based largely on photography and cinematography.2. Swimming inP. montereyensis involves the alternating antagonistic action of the subumbrellar swimming muscles and the elastic mesoglea.3. The swimming muscle consists of striated contractile elements arranged circularly in four discontinuous subumbrellar sheets and a sheet on the subumbrellar side of the velum. There is also a sheet of radially arranged fibers on the exumbrellar side of the velum. The four subumbrellar sheets are anchored to the bell along the perand interradii.4. The mesogleal skeleton consists of five components: (a) the matrix of the bell mesoglea, (b) optically visible fibers that traverse the bell from gastrodermal lamella to exumbrella, (c) the basement membrane or supporting lamella, (d) a system of joints, and (e) the velar mesoglea.5. The morphology, orientation, and distribution of the mesogleal fibers suggest that their major role is maintaining the radial integrity of the bell during deformation. The amount of stretch in a region of the bell wall during contraction is inversely proportional to the number of fibers per unit area there. In regions of the bell which are not deformed during contraction fibers are sparse or absent.6. Mesogleal volume remains constant during swimming. Locally the mesoglea is subjected to forces of stretch and compression, but the critical element in narrowing the bell involves bending or folding the mesoglea around a series of structural joints. The fulcrum of these joints is anchored to the exumbrella by concentrations of mesogleal fibers. The joints consist of eight adradial regions of highly deformable mesoglea lacking visible fibers. The regions are triangular in cross section and are separated from the remainder of the mesoglea (98–99 % of the total) by the gastrodermal lamella. A circular apical joint is also present.7. Sequential changes in shape and position of the bell relative to a fixed grid during contraction and recovery were measured in order to determine such parameters of swimming as rate of contraction, rate of expulsion of water, change in bell velocity during contraction and recovery, momentum, etc.8. The function of the velum was determined by cinematographic analysis of swimming animals both before and after removal of the velum. In normal swimming the velum serves mainly to constrict the aperture of the bell, thus increasing the velocity of expelled water, and hence increasing the force driving the medusa foward. Medusae swam with a greatly decreased velocity after velum removal.9. Turning is accomplished primarily by asymmetrical contraction of the exumbrellar velar radial muscles, whereby the velar aperture is displaced to one side, water is expelled obliquely, and the bell turns toward that same side. The ability to turn was lost after velum removal.10. Studies of the relationship between individual size and the various parameters of swimming inP. montereyensis show that: (a) the duration of the contraction phase of the swimming beat is roughly proportional to the square root of the subumbrellar circumference (or bell height); (b) smaller individuals swim faster relative to their bell height than do larger ones; (c) the velum is relatively better developed in small animals and plays a proportionately more important role during swimming.

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Struktur und Funktion des lokomotorischen Systems vonPolyorchis montereyensis (Cnidaria, Hydrozoa)

Kurzfassung Mit Hilfe histologischer und kinematographischer Methoden wurden Bau und Funktion des lokomotorischen Systems der AnthomedusePolyorchis montereyensisSkogsberg analysiert. Die Schwimmbewegungen resultieren aus der antagonistischen Wirkung der Muskulatur der Subumbrella und der elastischen Mesogloea. Struktur, Anordnung, Verteilung und Verankerung der Muskelzellen werden beschrieben. Die Körperschicht der Mesogloea besteht aus 5 Komponenten mit Skelettfunktion: der Matrix der Schirmmesogloea, einem System von Muskelfasern, der Stützlamelle, acht adradialen Verbindungssträngen von stark deformierbarer, fibrillenloser Mesogloea und der Mesogloea des Velums. Die Fribrillen und Verbindungsstränge der Mesogloea bewirken, daß die Schirmglocke während der Kontraktion deformiert werden kann. Die Funktion des Velums sowie die kontinuierlichen Veränderungen der Glocke in bezug auf Form und Lage während des Schwimmvorgangs, insbesondere die Geschwindigkeit der Kontraktion, des Wasserausstoßes und der Fortbewegung bei Individuen verschiedener Größe, wurden eingehend untersucht. Kleinere Medusen schwimmen relativ schneller als größere, was hauptsächlich auf stärkere Kontraktionen des Velums zurückzuführen ist.

     

Induction of metamorphosis from the larval to the polyp stage is similar in Hydrozoa and a subgroup of Scyphozoa (Cnidaria, Semaeostomeae)

Larvae of cnidarians need an external cue for metamorphosis to start. The larvae of various hydrozoa, in particular of Hydractinia echinata, respond to Cs⁺, Li⁺, NH₄ ⁺ and seawater in which the concentration of Mg²⁺ ions is reduced. They further respond to the phorbolester, tetradecanoyl-phorbol-13-acetate (TPA) and the diacylglycerol (DAG) diC8, which both are argued to stimulate a protein kinase C. The only well-studied scyphozoa, Cassiopea spp., respond differently, i.e. to TPA and diC8 only. We found that larvae of the scyphozoa Aurelia aurita, Chrysaora hysoscella and Cyanea lamarckii respond to all the compounds mentioned. Trigonelline (N-methylnicotinic acid), a metamorphosis inhibitor found in Hydractinia larvae, is assumed to act by delivering a methyl group for transmethylation processes antagonising metamorphosis induction in Chrysaora hysoscella and Cyanea lamarckii. The three species tested are scyphozoa belonging to the subgroup of semaeostomeae, while Cassiopea spp. belong to the rhizostomeae. The results obtained may contribute to the discussion concerning the evolution of cnidarians and may help to clarify whether the way metamorphosis can be induced in rhizostomeae as a whole is different from that in hydrozoa and those scyphozoa belonging to the subgroup semaeostomeae. Electronic Publication     

Coordinated modulation of cellular signaling through ligand-gated ion channels in Hydra vulgaris (Cnidaria, Hydrozoa)

Cnidarians lack well developed organs, but they have evolved the molecular and cellular components needed to assemble a nervous system. The apparent 'simplicity' of the cnidarian nervous net does not occur at the cellular level, but rather in the organisation of conducting systems. Cnidarian neurons are in fact electrically excitable, show the typical extended morphology and are connected by chemical synapses or gap junctions. They have been regarded as peptidergic, given the wealth of neuropeptides generally distributed along neurites and in cell bodies, supporting the hypothesis of a modulatory role in neurotransmission. However, the presence of clear-cored, as well as dense-cored synaptic vesicles in cnidarian neurons suggests both fast and slow synaptic transmission mechanisms. In fact, biochemical and functional evidence indicates that classical neurotransmitters and their metabolic partners are present in cnidarian tissues, where they are involved in coordinating motility and behavior. We have identified and characterized in Hydra tissues receptors to the inhibitory and excitatory amino acid neurotransmitters, GABA, glycine and NMDA, that are similar to mammalian ionotropic receptors in terms of their biochemical and pharmacological properties. These receptors appear to regulate pacemaker activities and their physiological correlates; in the live animal, they also affect feeding behavior, namely the duration and termination of the response elicited by reduced glutathione, with opposite actions of GABA and glycine or NMDA, respectively. These results suggest that modulation of cellular signaling through ligand-gated-ion channels is an ancient characteristic in the animal kingdom, and that the pharmacological properties of these receptors have been highly conserved during evolution.     

Distribution, ecology and morphology of Lytocarpia myriophyllum (Cnidaria: Hydrozoa), a Mediterranean Sea habitat former to protect

In the Mediterranean Sea, as well as in other parts of the word, intense bottom trawling threatens deep and mesophotic assemblages, compromising mainly the survivorship of erect organisms and of the habitat complexity they shape. Protection of species able to affect their habitats, by increasing spatial complexity and enhancing interspecific interactions, is crucial for biodiversity conservation. It is urgent to highlight the occurrence of those species which act as ecosystem engineers and/or habitat former to enhance awareness on their ecological role and to develop focused conservation strategies. Lytocarpia myriophyllum is the largest Leptomedusan hydroid of the Mediterranean Sea, with colonies up to 1 m high, and the most abundant Aglaopheniid in the eastern part of the North Atlantic Ocean. This species creates wide forests on soft bottoms stabilizing sediments, providing refuge and food for several other associated organisms and could be defined both a habitat former and an ecosystem engineer. Thanks to trimix diving here we report on new insights on the morphological, biological and ecological features of L. myriophyllum meadows from the Mediterranean Sea furnishing a baseline for protection plans focused on these facies. This work demonstrates that direct studies of mesophotic habitats allow to collect far more detailed information than grabs, ROVs, or towed camera arrays and highlights the urgent need to redefine the vertical extension of several marine protected areas.     

Growth and reproduction as a function of temperature and salinity inClava multicornis (Cnidaria,Hydrozoa)

Summary 1. Rates of growth (length increase of stolons) and of asexual reproduction (increase in number of polyps) were determined in secondaryClava multicornis colonies of a clone exposed to 12 different combinations of water temperature and salinity (12°, 17°, 22° C; 16 , 24 , 32 , 40 S). Sexual reproduction (via gonophores) has been observed only at 12° and 17° C; temperature and salinity ranges are narrower for sexual than for asexual reproduction.2. The data obtained are insufficient for a detailed analysis; they provide, however, interesting insights into the variability of growth and reproduction ofC. multicornis caused by different intensities of temperature and salinity.3. It appears that temperature requirements for maximum colony increase are reduced as the colony grows older.4. One feeding period per 24 hours seems insufficient for maximum growth and reproduction at the higher temperature levels, especially at 22° C.5. The different degrees of environmental stress endured during the initial period of transfer into the test combinations of temperature and salinity have affected the resulting colony size at least up to an age of 39 days. More appropriate criteria for assessment of rates of growth and reproduction are therefore the doubling times (number of days within which stolon length and polyp numbers taken 20 days after initiation of experiments have doubled).6. On the basis of doubling time values, increase in stolon length is progressively reduced with increasing water temperature (12°, 17°, 22° C). At 12° and 17° C stolons grow fastest in 32 , followed by 24 , 16 and 40 S; at 22° C stolon growth rates are identical in 32 and 24 S.7. Doubling times of polyp numbers per colony show a less obvious trend. In 56-day-old colonies, however, stolon length and polyp number are modified to similar degrees by the various temperatures and salinities offered. The sequence of temperatures causing fastest increase in polyp number is 12°>17°>22° C; the respective sequence of salinities reads: 24 , 32 , 16 , 40 S.8. Stolon length and polyp number per colony increase exponentially; most curves obtained exhibit undulations indicating endogenous growth rhythms.9. During the initial period of transfer into the final test media, asexual reproduction via budding seems to have been stimulated by a reduction in salinity.10. The doubling times obtained forC. multicornis are considerably longer than those found forCordylophora caspia and indicate that our culture conditions may have been suboptimal.

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Wachstum und Reproduktion als Funktion von Temperatur und Salzgehalt beiClava multicornis (Cnidaria, Hydrozoa)

Kurzfassung Einzelpolypen eines Klons vonC. multicornis Forskål wurden schrittweise in 12 verschiedene Temperatur-Salzgehalts-Kombinationen überführt und — während sie zu neuen Kolonien heranwuchsen — das Längenwachstum ihrer Stolonen, die Geschwindigkeit ihrer asexuellen Vermehrung durch Knospung neuer Hydranthen sowie die Gonophorenausbildung (sexuelle Fortpflanzung) registriert. Die erhaltenen Daten sind unzureichend für eine detaillierte Analyse, gewähren jedoch interessante Einblicke in die Bedeutung der verschiedenen Temperatur- und Salzgehaltsbedingungen für Wachstum und Vermehrung. Die anfängliche, schrittweise Überführung in die Testmedien verursacht per se Leistungsunterschiede, deren Auswirkungen sich mindestens bis zu einem Alter von 39 Tagen verfolgen lassen. Doubling times stellen daher objektivere Kriterien dar als absolute Zuwachswerte. Die doubling times von Kolonien, welche länger als 20 Tage in den Testmedien gewachsen waren, zeigen eine Verringerung der Stolonenzuwachsrate mit steigender Temperatur (12°, 17°, 22° C). Die Reihenfolge der fördernden Wirkung der einzelnen Salzgehaltsstufen ergibt sich zu 32 , 24 , 16 , 40 S. Im Prinzip ähnliche Verhältnisse liegen hinsichtlich der asexuellen Vermehrungsrate vor. Bemessen an den getesteten Kriterien scheinen die Temperaturansprüche mit zunehmendem Koloniealter abzunehmen. Die errechneten doubling times sind wesentlich länger als beiCordylophora; möglicherweise deutet dieser Unterschied auf inadäquate Kulturbedingungen (Fütterung, Wasserbewegung) hin.

     

Phylogenetics and evolution of Capitata (Cnidaria: Hydrozoa), and the systematics of Corynidae

Nawrocki, A. M., Schuchert, P. & Cartwright, P. (2009). Phylogenetics and evolution of Capitata (Cnidaria: Hydrozoa), and the systematics of Corynidae.—Zoologica Scripta, 39, 290–304. Generic‐ and family level classifications in Hydrozoa have been historically problematic due to limited morphological characters for phylogenetic analyses and thus taxonomy, as well as disagreement over the relative importance of polyp vs. medusa characters. Within the recently redefined suborder Capitata (Cnidaria: Hydrozoa: Hydroidolina), which includes 15 families and almost 200 valid species, family level relationships based on morphology alone have proven elusive, and there exist numerous conflicting proposals for the relationships of component species. Relationships within the speciose capitate family Corynidae also remain uncertain, for similar reasons. Here, we combine mitochondrial 16S, and nuclear 18S and 28S sequences from capitate hydrozoans representing 12 of the 15 valid capitate families, to examine family level relationships within Capitata. We further sample densely within Corynidae to investigate the validity of several generic‐level classification schemes that rely heavily on the presence/absence of a medusa, a character that has been questioned for its utility in generic‐level classification. We recover largely congruent tree topologies from all three markers, with 28S and the combined dataset providing the most resolution. Our study confirms the monophyly of the redefined Capitata, and provides resolution for family level relationships of most sampled families within the suborder. These analyses reveal Corynidae as paraphyletic and suggest that the limits of the family have been underestimated. Our results contradict all available generic‐level classification schemes for Corynidae. As classification schemes for this family have been largely based on reproductive characters such as the presence/absence of a medusa, our results suggest that these are not valid generic‐level characters for the clade. We suggest a new taxonomic structure for the lineage that includes all members of the newly redefined Corynidae, based on molecular and morphological synapomorphies for recovered clades within the group.     

Different roles of GABA and glycine in the modulation of chemosensory responses in Hydra vulgaris (Cnidaria,Hydrozoa)

Phylogenetic studies suggest that GABA and glycine receptors derive, as a result of divergent evolution, from a common ancestral protoreceptor originated in a unicellular organism. This raises the possibility that members of the ligand-gated ion channels (LGIC) superfamily might be widely present in living organisms including bacteria and primitive invertebrates. High-affinity GABA receptors occur in the tissues of Hydra vulgaris whose pharmacological characteristics compare with those of mammalian ionotropic GABA receptors. Behavioural studies have shown that activation of these GABA _A -like receptors by their allosteric modulators increases the duration of response to reduced glutathione (GSH). Recently, strychnine-sensitive glycine receptors have been shown to occur in Hydra tissues. Activation of these glyR also results in increased duration of the response to GSH. In order to investigate the contribution of endogenous transmitters to the modulation of the feeding response, we studied the effects of exposing the polyps to brief depolarizing pulses prior to the GSH test. A severe inhibition of the response was observed following exposure to KCl or veratridine. Administration of GABA or muscimol counteracted the effects of the pulses in a dose-dependent manner. The effects of GABA or muscimol were suppressed by the GABA _A -specific antagonist gabazine both in pulse-untreated and treated polyps. By contrast, glycine and its agonist taurine were not able to restore the physiological duration of response in pulse-treated Hydra, while another glyR agonist, β-alanine, partially reduced the pulse-induced inhibition. We conclude that GABA appears to be the major inhibitory transmitter responsible for the regulation of the feeding response. Molecular studies aimed at identifying GABA receptor subunits are in progress.     

Species in the genus Turritopsis (Cnidaria, Hydrozoa): a molecular evaluation

Mitochondrial ribosomal gene sequences were used to investigate the status of several populations of hydromedusae belonging to the genus Turritopsis (family Oceaniidae). Several nominal species have been described for this genus, but most of them had been synonymized and attributed to one cosmopolitan species, Turritopsis nutricula . A recent revision based on morphological and reproductive characters, however, has shown that many different populations can be distinguished and that several of the nominal Turritopsis species are likely valid biological species. Our investigation using molecular sequence data of 16S mitochondrial gene confirms these results. The Mediterranean Turritopsis must be attributed to Turritopsis dohrnii and the Turritopsis of New Zealand must be referred to Turritopsis rubra . The situation of the Japanese Turritopsis is more complex, though all sampled populations are clearly distinct from T. nutricula , a species likely confined to the Western Atlantic. The Japanese Turritopsis fall into three widely separated lineages. One of them, corresponding likely to Turritopsis pacifica , is closely related to T. rubra . A second clade, which potentially represents an as yet undescribed species, produces smaller medusae than T. pacifica and is morphologically distinguishable from it. Finally, a third group was distinguished by a single haplotype sequence that is identical with a Mediterranean sample of T. dohrnii . It is postulated that the last group of Japanese Turritopsis is likely a recent introduction, most probably by human activity. A survey of all known and potentially valid Turritopsis species is given in table format to facilitate identifications and future revisory work.     

Hydranth structure and digestion rate as a function of temperature and salinity inClava multicornis (Cnidaria,Hydrozoa)

Summary 1. Responses to 12 different combinations of constant levels of temperature and salinity were tested in the colonial athecate hydroidClava multicornis. Criteria measured were: (a) length and width of hydranth bodies, (b) number and length of tentacles and (c) rate of digestion.2. Test colonies were obtained by allowing single hydranths — cut off from an individual primary colony — to regenerate via asexual reproduction into new secondary colonies.3. In the resulting — genetically identical — material, all criteria tested vary as a function of the different environments offered.4. In 16, 24 and 32 S hydranth length reaches maximum values at 12°C, followed by 17° and then 22°C. With increasing salinity, hydranth length declines markedly at 12°C, while there is little or no decline at 17°C and a definite increase from 24 to 32 S at 22°C.5. Hydranth width varies less extensively; in general, it follows similar patterns as does hydranth length.6. Tentacle number per hydranth tends to be positively related to the size of the hydranth body; it decreases with increasing temperature. Salinity levels producing maximum tentacle numbers vary with temperature; maxima are found at 12°C in 16, at 17°C in 24 and at 22°C at 32 S.7. Tentacle length — although a more variable criterion — is affected similarly to tentacle number; it attains, however, relatively higher values at 17°C.8. In regard to the hydranth dimensions measured, combinations of low temperatures/low salinities and of high temperatures/high salinities tend to produce maximum values.9. Rate of digestion is taken here to be identical to the time elapsing between completed food intake and defecation. This time span is reduced with increasing temperature. In all temperature levels, digestion time is shortest in 32 S.10. The results presented above compare well with those obtained earlier under similar conditions onCordylophora caspia.11. It is assumed that the structural modifications of hydranths affect rate and efficiency of exchanges between organism and environment and may thus represent a means of metabolic adjustment.

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Hydranthenstruktur und Verdauungsrate als Funktion von Temperatur und Salzgehalt beiClava multicornis (Cnidaria, Hydrozoa)

Kurzfassung Erbgleiche Polypen vonClava multicornis Forskål wurden in 12 verschiedenen Temperatur-Salzgehalts-Kombinationen durch asexuelle Vermehrung zu neuen Kolonien herangezogen und an den adulten Hydranthen dieser erbgleichen Sekundärkolonien Länge und Breite des Hydranthenkörpers, Anzahl und Länge der Tentakel sowie die Verdauungsgeschwindigkeit ermittelt. Alle gemessenen Parameter verändern sich — zum Teil sogar erheblich — als Funktion von Temperatur und Salzgehalt. Die Veränderungen der Hydranthendimensionen führen zu Verschiebungen der Oberfläche-Volumen-Quotienten. Da der Stoffaustausch zwischen Kolonie und Umwelt primär im Bereich der Hydranthen erfolgt, kommt diesen Verschiebungen vermutlich eine Bedeutung zu im Rahmen der Stresskompensation (nicht-genetische Adaptation). Die erzielten Ergebnisse bestätigen frühere Untersuchungen vonKinne an dem naheverwandten HydroidpolypenCordylophora caspia.

     

First identification and localization of a visual pigment in Hydra (Cnidaria, Hydrozoa)

The cnidarian Hydra does not possess identified photoreceptive structures or specialized cells for light detection; nevertheless, it shows a marked photosensitivity. So far no evidence has been previously reported about the localization of the proteins involved in the photoresponse. We used polyclonal antibodies and immunofluorescence microscopy on whole-mount Hydra to identify a putative rhodopsin-like protein. Our results show an immunoreactivity in the ectodermal layer of Hydra, which corresponds in position to the nervous epidermal sensory cells. These data provide the first identification of a rhodopsin-like protein in a phylogenetically old invertebrate and give a new insight into the Hydra photoreceptive response.     

Redescription of Licnophora chattoni Villeneuve-Brachon, 1939 (Ciliophora, Spirotrichea), associated with Zyzzyzus warreni Calder, 1988 (Cnidaria, Hydrozoa)

Licnophora chattoni, found in association with Zyzzyzus warreni, a tubulariid hydroid epizoic in sponges from São Sebastião (SP, Brazil), is redescribed and illustrated using light and electron microscopy. The ciliate has a flexible, transparent body formed by an oval anterior region linked to the posterior basal disc via a flexible neck region. Numerous cortical granules are observed scattered throughout the body and densely packed along the neck. The adoral zone is formed by about 81 external and 24 infundibular paramembranelles. The paroral membrane, formed by a row of long cilia arranged in monokinetids, extends through a groove in the body to the adhesive disc. Two dorsal kinetids are present along the right body margin and around the neck. The adhesive disc (18 μm in diameter) lacks cilia in the area above the velum. The velum covers a row of dikinetids bearing long cilia and four dikineties, two or three of which are interrupted on the ventral surface. Nine to twelve macronuclear nodules connected by isthmuses are distributed in the cytoplasm, plus two nodules located in the adhesive disc and between those there is an ovate micronucleus.     

Epiphytic hydroids (Cnidaria,Hydrozoa) contribute to a higher abundance of caprellid amphipods (Crustacea,Peracarida) on macroalgae

The behaviour of returning Salmo salar (Linnaeus, 1758) approaching, and attempting to pass low-head weirs remains relatively unknown. A radio telemetry array was created at a low-head weir to enable the behaviour of S. salar (n = 120) to be observed as they approached and attempted to pass the barrier. The majority of fish successfully passed the barrier on their first or second attempt, some individuals required 11 attempts prior to successful passage occurring. Mean delay at the barrier per fish was 47.8 h (±SD 132.0 h), range 15 min to 31 days. Passage success on a fish’s initial attempt was significantly predicted by the amount of searching a fish undertook, fork length, and fat content. Fish were more likely to have a successful first passage attempt if it was smaller with a low fat content and exerted a greater effort in searching for a passage channel. Small-scale barriers cause delays and increased energy expenditure in migrating fish. Barriers may be creating an anthropogenic selection pressure for traits which enable passage success. The impact of a delay at a barrier and increased energy expenditure on reproduction and gonad development remains unknown but is likely to be negatively impacted by instream anthropogenic structures.     

Post-embryonic larval development and metamorphosis of the hydroidEudendrium racemosum (Cavolini) (Hydrozoa,Cnidaria)

The morphology and histology of the planula larva ofEudendrium racemosum (Cavolini) and its metamorphosis into the primary polyp are described from light microscopic observations. The planula hatches as a differentiated gastrula. During the lecithotrophic larval period, large ectodermal mucous cells, embedded between epitheliomuscular cells, secrete a sticky slime. Two granulated cell types occur in the ectoderm that are interpreted as secretory and sensorynervous cells, but might also be representatives of only one cell type with a multiple function. The entoderm consists of yolk-storing gastrodermal cells, digestive gland cells, interstitial cells, cnidoblasts, and premature cnidocytes. The larva starts metamorphosis by affixing its blunt aboral pole to a substratum. While the planula flattens down, the mucous cells penetrate the mesolamella and migrate through the entoderm into the gastral cavity where they are lysed. Subsequently, interstitial cells, cnidoblasts, and premature cnidocytes migrate in the opposite direction, i.e. from entoderm to ectoderm. Then, the polypoid body organization, comprising head (hydranth), stem and foot, all covered by peridermal secretion, becomes recognisable. An oral constriction divides the hypostomal portion of the gastral cavity from the stomachic portion. Within the hypostomal entoderm, cells containing secretory granules differentiate. Following growth and the multiplication of tentacles, the head periderm disappears. A ring of gland cells differentiates at the hydranth's base. The positioning of cnidae in the tentacle ectoderm, penetration of the mouth opening and the multiplication of digestive gland cells enable the polyp to change from lecithotrophic to planktotrophic nutrition.     

Ecological and morphological characteristics of Ephelota gemmipara (Ciliophora,Suctoria), epibiontic on Eudendrium racemosum (Cnidaria,Hydrozoa) from the Adriatic Sea

The relationship between the suctorian Ephelota gemmipara and the large hydroid Eudendrium racemosum from the North Adriatic Sea has been studied over its full annual cycle. Ephelota gemmipara settles on the perisarc of the hydroid, usually close to the hydranths in order to exploit the hydroid's food discharges. The life cycle of E. gemmipara is influenced by temperature variations and by its relationship with the host. The hydroid shows an active phase in the summer, and it gets through the adverse winter season forming resting stages. In April, when temperature increases, the hydroid starts its active phase and it is colonized by suctorians. From May to September the suctorians produce multiple buds (swarmers) that detach from the parental cells to settle on an Eudendrium colony. The abundance of the suctorian peaks in September, with more than 1.2 million ind. m^?2. Their proliferation coincides with the maximal abundance of their host and the highest water temperatures. On the contrary, sexual reproduction and the encystment occur when the temperature and the abundance of E. racemosum decrease. Lastly, we also report the presence of symbionts such as bacteria and the parasitic protozoans Tachyblaston ephelotensis and Enigmocoma acinetarum.     

Biodiversity of prokaryotic communities associated with the ectoderm of Ectopleura crocea (Cnidaria, Hydrozoa)

The surface of many marine organisms is colonized by complex communities of microbes, yet our understanding of the diversity and role of host-associated microbes is still limited. We investigated the association between Ectopleura crocea (a colonial hydroid distributed worldwide in temperate waters) and prokaryotic assemblages colonizing the hydranth surface. We used, for the first time on a marine hydroid, a combination of electron and epifluorescence microscopy and 16S rDNA tag pyrosequencing to investigate the associated prokaryotic diversity. Dense assemblages of prokaryotes were associated with the hydrant surface. Two microbial morphotypes were observed: one horseshoe-shaped and one fusiform, worm-like. These prokaryotes were observed on the hydrozoan epidermis, but not in the portions covered by the perisarcal exoskeleton, and their abundance was higher in March while decreased in late spring. Molecular analyses showed that assemblages were dominated by Bacteria rather than Archaea. Bacterial assemblages were highly diversified, with up to 113 genera and 570 Operational Taxonomic Units (OTUs), many of which were rare and contributed to <0.4%. The two most abundant OTUs, likely corresponding to the two morphotypes present on the epidermis, were distantly related to Comamonadaceae (genus Delftia) and to Flavobacteriaceae (genus Polaribacter). Epibiontic bacteria were found on E. crocea from different geographic areas but not in other hydroid species in the same areas, suggesting that the host-microbe association is species-specific. This is the first detailed report of bacteria living on the hydrozoan epidermis, and indeed the first study reporting bacteria associated with the epithelium of E. crocea. Our results provide a starting point for future studies aiming at clarifying the role of this peculiar hydrozoan-bacterial association.     

Four new species of Hydromedusae (Cnidaria, Hydrozoa) from the coast of south-western Africa

Four previously unknown species of hydromedusae are described from plankton samples collected during oceanographic cruises undertaken along the south-western coast of Africa. The Anthomedusa Bythotiara capensis sp.n., known from a single specimen, was found in shallow waters of the Cape region (South Africa). The Leptomedusa Margalefia intermedia gen. et sp.n. collected in Namibian waters, possesses desmoneme cnidocysts. This category of cnidocysts is previously unreported in the Leptomedusae, but is common in Anthomedusae and in the limnomedusan family Proboscidactylidae. Margalefia seems to occupy a position intermediate between the families Tirannidae and Laodiceidae. Two species of Limnomedusae, Aglauropsis edwardsii sp.n. and Proboscidactyla menoni sp.n. were frequent at several stations in Namibian waters between depths of 50 m and the surface.     

Trends in the Diversity,Distribution and Life History Strategy of Arctic Hydrozoa (Cnidaria)

This is the first attempt to compile a comprehensive and updated species list for Hydrozoa in the Arctic, encompassing both hydroid and medusa stages and including Siphonophorae. We address the hypothesis that the presence of a pelagic stage (holo- or meroplanktonic) was not necessary to successfully recolonize the Arctic by Hydrozoa after the Last Glacial Maximum. Presence-absence data of Hydrozoa in the Arctic were prepared on the basis of historical and present-day literature. The Arctic was divided into ecoregions. Species were grouped into distributional categories according to their worldwide occurrences. Each species was classified according to life history strategy. The similarity of species composition among regions was calculated with the Bray-Curtis index. Average and variation in taxonomic distinctness were used to measure diversity at the taxonomic level. A total of 268 species were recorded. Arctic-boreal species were the most common and dominated each studied region. Nineteen percent of species were restricted to the Arctic. There was a predominance of benthic species over holo- and meroplanktonic species. Arctic, Arctic-Boreal and Boreal species were mostly benthic, while widely distributed species more frequently possessed a pelagic stage. Our results support hypothesis that the presence of a pelagic stage (holo- or meroplanktonic) was not necessary to successfully recolonize the Arctic. The predominance of benthic Hydrozoa suggests that the Arctic could have been colonised after the Last Glacial Maximum by hydroids rafting on floating substrata or recolonising from glacial refugia.