Alteration of photoresponses involved in diel vertical migration of a crab larva by fish mucus and degradation products of mucopolysaccharides

Photoresponses involved in the descent phase of nocturnal diel vertical migration (DVM) of larvae of the crab Rhithropanopeus harrisii were measured in a laboratory system that mimicked the underwater angular light distribution. The test hypothesis was that kairomones from fish that activate zooplankton photoresponses involved in DVM are derived from polysaccharides from the external mucus of fishes. Studies considered fish mucus from the mummichog (Fundulus heteroclitus) and disaccharides (originating from chondroitin sulfate A and heparin polysaccharides) that are likely constituents of fish mucus. R. harrisii larvae descend at sunrise with an isolume and remain near the isolume during the day. Since depth maintenance near the isolume depends upon a negative phototaxis, the lowest light intensity (threshold) that induces this response was used to quantify the effects of the test chemicals. It was predicted that exposure to fish kairomones would lower the photoresponse threshold, thereby resulting in larvae remaining deeper in the water column where light for visual predation was reduced. The photoresponse threshold declined as the concentration of fish mucus increased. Disaccharides originating from chondroitin sulfate A and heparin also decreased the photoresponse threshold as compared to responses in aged, filtered seawater. Collectively, the results support the hypothesis and indicate that disaccharide degradation products of predator mucus containing sulfated and acetylated amines can serve as kairomones.     

Laboratory studies of predation by the ctenophore Mnemiopsis leidyi on the early stages in the life history of the bay anchovy, Anchoa mitchilli

The lobate ctenophore, Mnemiopsis leidyi, consumed eggs andlarvae of the bay anchovy, Anchoa mitchilli, in laboratory experiments.This ctenophore exhibited a type I functional response to increasesin egg densities without reaching saturation at high prey densities.Clearance rate increased with increasing experimental containervolume. There was a 3-fold increase in the volume of water clearedby 2.0–2.5-cm ctenophores and a >5-fold increase for4.5–5.0-cm ctenophores in 15–1 versus 100–200–1containers. Clearance rate was dependent on tenophore lengthbut was probably underestimated for the larger animals due tocontainer effects. The presence of various densities of alternateprey, Acartia hudsonica or Anemia sp. nauplii, in addition toAnchoa mitchilli eggs did not affect the clearance rates onthe eggs alone. Comparison of clearance rates of 2.0–2.5-cmctenophores on various ages of starved and fed bay anchovy larvaeindicated that predation may be higher on yolk-sac larvae thanon eggs but decreases as the larvae grow. After 3 days posthatch starved anchovy larvae become more vulnerable to predationthan fed larvae. The ctenophore, M.leidyi has the potentialto inflict substantial predation pressure on early stages inthe life history of bay anchovy. ¹ Present address: University of Maryland, Center for Environmentaland Estuarine Studies, Chesapeake Biological Laboratory, POBox 38, Solomons, MD 20688-0038, USA     

Rapid habituation by mosquito larvae to predator kairomones

Larvae of some species of mosquitoes have been shown to respond to water‐borne kairomones from predators by reducing bottom‐feeding and replacing it with surface filter‐feeding, which uses less movement and is thus less likely to attract a predator. However, if no predator attack takes place, then it would be more efficient to use a risk allocation strategy of habituating their response depending on the predator and the overall risk. The larvae of Culiseta longiareolata Macquart live in temporary rain‐filled pools, where they are exposed to a high level of predation. Within one hour, they responded to kairomones from dragonfly or damselfly nymphs, or to the fish Aphanius, by significantly reducing bottom‐feeding activity. Continued exposure to the predator kairomones resulted in habituation of their response to damselflies, a slower habituation to fish, but no habituation to dragonflies even after 30 h. In contrast, the larvae of Culex quinquefasciatus Say normally live in highly polluted and thus anaerobic water, where the predation risk will be much lower. They also showed a significant reduction in bottom‐feeding after 1 h of exposure to predator kairomones but had completely habituated this response within 6 h of continuous exposure. Some species of mosquito larvae can thus show a very rapid habituation to predator kairomones, while others only habituate slowly depending on the predator and overall predation risk.     

Bacterial communities associated with the ctenophores Mnemiopsis leidyi and Beroe ovata

Residing in a phylum of their own, ctenophores are gelatinous zooplankton that drift through the ocean's water column. Although ctenophores are known to be parasitized by a variety of eukaryotes, no studies have examined their bacterial associates. This study describes the bacterial communities associated with the lobate ctenophore Mnemiopsis leidyi and its natural predator Beroe ovata in Tampa Bay, Florida, USA. Investigations using terminal restriction fragment length polymorphism (T-RFLP) and cloning and sequencing of 16S rRNA genes demonstrated that ctenophore bacterial communities were distinct from the surrounding water. In addition, each ctenophore genus contained a unique microbiota. Ctenophore samples contained fewer bacterial operational taxonomic units (OTUs) by T-RFLP and lower diversity communities by 16S rRNA gene sequencing than the water column. Both ctenophore genera contained sequences related to bacteria previously described in marine invertebrates, and sequences similar to a sea anemone pathogen were abundant in B.?ovata. Temporal sampling revealed that the ctenophore-associated bacterial communities varied over time, with no single OTU detected at all time points. This is the first report of distinct and dynamic bacterial communities associated with ctenophores, suggesting that these microbial consortia may play important roles in ctenophore ecology. Future work needs to elucidate the functional roles and mode of acquisition of these bacteria.     

Growth dynamics of a ctenophore (Mnemiopsis) in relation to variable food supply. II. Carbon budgets and growth model

Our goal was to test our understanding of ingestion, assimilationefficiency and metabolism for Mnemiopsis mccradyi by formulatingand validating a simulation model of growth under differentconditions of food availability. The model was based on a carbonbudget approach using formulations derived from empirical results,including how each process was affected by food availabilityand ctenophore size. An experimentally measured carbon budgetfor pulsed food availability indicated that, relative to totalingestion, growth was high (17–48%), respiration plusorganic release was relatively low (24–48%) and little(<10%) of the ingested carbon was unaccounted for. New laboratoryinvestigations of feeding and assimilation efficiency were necessaryto refine the formulations so that model predictions comparedfavorably with a variety of laboratory measurements of growth,and growth efficiency, as well as the complete experimentallymeasured carbon budget. The refined model predicted a high ratioof growth to metabolism (>2) and a high gross growth efficiency(>30%) for smaller ctenophores at high food concentrations(>20 prey l^–1). Both growth rates and growth efficiencieswere predicted to decrease for larger ctenophores. Model predictionswere generally consistent with experimental results, includinginvestigations using pulsed food availability to simulate environmentalpatchiness. Although the model underpredicted ctenophore growthin some experiments at low food densities, the model predictionof a minimum prey concentration of about 8 l^–1 (24 µgC l^–1) for sustaining a ctenophore population of reproductivesize agreed with field observations.     

Benthic ctenophores (Platyctenida: Coeloplanidae) in South Florida: predator–prey interactions

The primary goal of this study was to demonstrate, from field observations and laboratory experiments, some key trophic roles of benthic ctenophores as predators and prey in subtropical communities. We examined individuals of two benthic platyctenid species: Coeloplana waltoni, a minute epibiont on octocorals in exposed, open‐water settings; and Vallicula multiformis, an associate of calm‐water biofouling communities and floating Sargassum spp. Laboratory observations of individuals of both ctenophore species revealed frequent capture and ingestion of diverse zooplankton taxa, especially crustaceans. Laboratory predation trials demonstrated the capture of dolphinfish (Coryphaena hippurus) eggs and larvae by both ctenophore species. Dolphinfish eggs and larvae larger than individuals of C. waltoni were captured but not ingested during 2‐h trial periods. These prey items were sometimes purloined and ingested by polyps of the ctenophore's octocoral host. Ingestion of dolphinfish eggs and larvae by individuals of C. waltoni was observed, however, after longer periods of exposure to prey. In predation trials, dolphinfish eggs and larvae were both captured and ingested by larger individuals of the ctenophore species V. multiformis. Field and laboratory observations revealed diverse invertebrate and fish taxa that prey on both ctenophore species. In the laboratory, the mean daily per capita consumption of individuals of C. waltoni by a pomacanthid fish ranged 0.5–2.8 individuals, and ranged 2.6–3.6 individuals for predation by an ovulid mollusc. Field population densities of these predators ranged 0.1–0.7 individuals per m² for the pomacanthid, and 0.2–1.1 individuals per m² for the mollusc. Laboratory feeding observations demonstrated frequent consumption of individuals of V. multiformis by a sea anemone, and by three species of brachyuran crabs. Field observations revealed eight fishes that probably feed incidentally on individuals of V. multiformis. These findings add to the limited knowledge base of predator–prey dynamics in both C. waltoni and V. multiformis.     

Comparative experimental studies on the feeding of chaetognaths and ctenophores

There is very little quantitative experimental information onfeeding in chaetognaths and ctenophores because of the difficultyof obtaining undamaged specimens. As important plankton predators,a knowledge of their daily rations as a function of food concentrationis vital to ecosystem models of water column trophodynamics.At specified food concentrations, three ctenophore species alwaysingested higher specific daily rations than three species ofchaetognaths. Food consumption of ctenophores increased linearlythroughout the food concentration range (exceeding 10⁶ µgC/m³ Although food consumption of chaetognaths reached an upperlimit at very high food concentrations, it seemed unlikely thatsatiation would occur under environmental conditions. Below10,000 food items/m³ chaetognaths were unable to obtain l% oftheir specific daily ration, although the few environmentalstudies based on gut contents suggest substantial feeding evenbelow this concentration. The apparent threshold may reflectthe environmental patch density necessary for feeding to begin.At high food concentrations daily rations of newly-hatched ctenophoresand chaetognaths can exceed well over 10 times their body carbon.In ctenophores even adults can consume these extremely largerations although chaetognaths follow the more usual pattern,where ration decreases throughout life. Gut residence time offood (where single items were ingested) increased with age andsize of food item selected in Sagitta hispida from one to twohours, levelling off after animals reached 6 mm. Mnemiopsismccradyi maintained a uniform digestion time of one hour asit grew, except for a brief period when it first selected larger-sizedfood. In older animals of both species, multiple ingestion becamemore common, where gut residence time was directly proportionalto number of food items consumed, ranging up to 5 hours forfive or more food items. Chaetognaths showed more variabilityof gut residence time than did ctenophores. It was concludedthat these animals are much better at catching food in the environmentthan laboratory experiments would suggest.     

Activation of brine shrimp nauplii photoresponses involved in diel vertical migration by chemical cues from visual and non-visual planktivores

The effects of exposure to visual and non-visual planktivoreson the photoresponses involved in the descent phase of nocturnaldiel vertical migration (DVM) of bnne shrimp (Anemia franciscana)naupliar larvae were measured in a laboratory system that mimickedthe underwater angular light distribution. This species wasused as a model for testing the general effects of differentplanktivores because a previous study demonstrated that naupliarphotoresponses were activated by exposure to one fish speciesthat was a visual planktivore, but did not co-exist with brineshrimp. The present study tested other fish and non-visual planktivores(ctenophores, chaetognaths, blue crab postlarvae). Photoresponseswere activated by 1 day exposure to: (i) three species of fish(Atlantic menhaden larvae, mummichog and pinfish) and (ii) waterthat had previously contained the fish or ctenophores. Thus,chemical cues from both visually and non-visually hunting planktivoresactivated photoresponses, contrary to the hypothesis that nocturnalDVM functions for avoidance of visual planktivores. Activationoccurred within 5 min, indicating that brine shrimp naupliihave a phenotypic response to zooplankton planktivores. Photoresponsesensitivity decreased with decreasing concentration of chemicalcue, indicating that activation of DVM should vary with planktivoreabundance. In contrast, photoresponse activation was very weakafter exposure to the physical presence of two non-visual, verticallymigrating planktivores (blue crab postlarvae and chaetognaths).The results support the predictions that zoo-plankton DVM patternshould vary with exposure to different planktivore types andthat migration amplitude should increase with increasing planktivoreabundance.     

Growth dynamics of a ctenophore (Mnemiopsis) in relation to variable food supply. I. Carbon biomass, feeding, egg production, growth and assimilation efficiency

This paper contains new experimental data on the growth dynamicsof a lobate coastal ctenophore, Mnenmiopsis mccradyi, whichadd significantly to our understanding of the nutritional ecologyof ctenophores and their role as opportunistic predators. Theseexperimental observations were necessary to refine the dynamiccarbon budget presented as a simulation model in another report.The ratio of carbon biomass to dry weight may vary several-folddepending on the nutritional state and size from >12% inwell-fed larvae to <1% in starved adults. Feeding effort(clearance rate) is higher for previously starved animals, fallingsharply within a few hours after re-exposure to food. Directvisual observations of feeding activity of animals confirmedthis. Assimilation efficiency can be high (72%) in these animalsbut if they continue to feed at high food concentrations, incomingfood displaces material which is only partially digested andassimilation efficiency decreases substantially. Except at verylow food concentrations, growth efficiency ranges between 20and 45%. Mnemiopsis, begins to produce eggs at a size much lessthan its maximum. Egg production is very sensitive to food supply,and somatic growth is favored over egg production at low fooddensities. Even though several thousand eggs may be producedover a few days, they represent <2% day^–1 of the carbonbiomass of the ctenophore and several-fold less than respiratorycarbon.     

Ecological and developmental dynamics of a host-parasite system involving a sea anemone and two ctenophores

The lined sea anemone Edwardsiella lineata has evolved a derived parasitic life history that includes a novel body plan adapted for life inside its ctenophore hosts. Reputedly its sole host is the sea walnut, Mnemiopsis leidyi, a voracious planktivore and a seasonally abundant member of many pelagic ecosystems. However, we have observed substantially higher E. lineata prevalence in a second ctenophore species, the ctenophore predator Bero? ovata. The interplay among these 3 species has important conservation consequences as M. leidyi introductions are thought to be responsible for the severe depletion of numerous commercial fisheries in the Mediterranean basin, and both E. lineata and B. ovata have been proposed as biological controls for invasive M. leidyi. Over a 3-yr period (2004-2006), we collected 8,253 ctenophores from Woods Hole, Massachusetts, including M. leidyi, B. ovata, and a third ctenophore, Pleurobrachia pileus, and we recorded E. lineata infection frequencies, parasite load, and parasite location. We also conducted laboratory experiments to determine the likely mechanisms for parasite introduction and the effect of each host on parasite development. We observed peak E. lineata infection frequencies of 0% in P. pileus, 59% in M. leidyi, and 100% in B. ovata, suggesting that B. ovata could be an important natural host for E. lineata. However, in laboratory experiments, E. lineata larvae proved far more successful at infecting M. leidyi than B. ovata, and E. lineata parasites excised from M. leidyi exhibited greater developmental competence than parasites excised from B. ovata. Although we show that E. lineata is efficiently transferred from M. leidyi to B. ovata when the latter preys upon the former, we conclude that E. lineata larvae are not well adapted for parasitizing the latter species and that the E. lineata parasite is not well adapted for feeding in B. ovata; these developmental and ecological factors underlie the host specificity of this recently evolved parasite.     

Peptide analogs of the mud crab pumping pheromone: structure-function studies

In the mud crab, Rhithropanopeus harrisii (Gould), larval releaseis highly synchronous and is controlled by peptide pheromonesreleased from the hatching eggs. The pheromones and syntheticpeptide analogs, containing neutral amino acids at the arnino-terminusand a basic amino acid at the carboxy-terminus, evoke larvalrelease or pumping behavior in the female. Here we describestructure–function relationships between behavioral responsesand peptide analogs of the pumping pheromone. Compounds testedincluded 11 peptides containing 1–7 neutral amino acidresidues and a carboxy-terminal arginine residue, and modifiedarginine compounds. Complex feeding-stimulant mixtures and groupsof peptides were tested to provide information about the numberof different types of receptors. Pumping behavior was describedin terms of response threshold, maximum percentage responseand effective concentration range. All carboxy-terminal argininepeptides tested evoked pumping responses. Response thresholdsranged from 10^-15 M for the most potent peptide Ile-Gly-Argto 10^-8 M for the least potent Tyr-Arg. Effective concentrationranges for the peptides varied from one to four orders of magnitudeand the responses normally declined with further increase inconcentration. Maximum percentage response was largely independentof response threshold and ranged from 30–60%, comparableto assay responses of larval hatch water. Responses within sizegroups of peptides were more similar than between groups. Studieswith modified arginines showed that hydrophobic structures similarin size and shape to small peptides evoked responses. Stericallyhindered hydrophobic moieties were ineffective. Tests with mixturesof active compounds support the hypothesis that there is onetype of pumping peptide receptor.     

Behavioral responses to fish kairomones and autotomy in a damselfly

The threat-sensitivity hypothesis predicts that prey species assess and adjust their behavior in accordance with the magnitude of the threat posed by a predator. A largely overlooked characteristic of a prey that will affect its sensitivity to predators is its history of autotomy. We studied threat-sensitive behavior to fish kairomones in larvae of Ischnura elegans damselflies, which had undergone autotomy, from a fishpond and from a fishless pond. In agreement with their higher perceived risk, larvae from the fishpond showed fewer rigid abdomen bends, foraged less and walked more slowly than larvae from the fishless pond. In line with their higher vulnerability to predators, larvae without lamellae spent less time foraging than larvae with lamellae. There was a decrease in swimming activity in the presence of fish kairomones except for larvae with lamellae from the fishless pond. This may reflect differences in vulnerability of larvae without lamellae between pond types. Such context-dependent responses in activity to kairomones should be kept in mind when evaluating the ability of a prey to recognize kairomones.     

Effects of UV-B radiation on near-surface zooplankton of puget sound

Summary An increase in incident solar ultraviolet irradiation, resulting from possible deterioration of the stratospheric ozone layer, would have important biological effects. Though the oceans are relatively opaque to UV radiation, compared to visible light, increases in incident UV may affect organisms living within the first few meters of the sea surface.Shrimp larvae, crab larvae, and euphausids were exposed to various low levels of simulated solar UV radiation (UV-B, 290–315 nm) under laboratory conditions. Comparisons between solar and artificial spectra were based on spectroradiometric measurements converted to erythemally effective irradiance. These zooplankton tolerated UV-B irradiance levels up to threshold levels with no significant reduction in survival or developmental rates compared to control organisms. Beyond the threshold levels, activity, development, and survival rapidly declined. The apparent UV thresholds are near present incident UV levels.Observed survival threshold levels for each experimental group were superimposed on seasonal solar incident UV levels at the experimental site. These threshold levels appeared to be exceeded by median ambient UV levels late in the season of surface occurrence of each species. UV increases resulting from ozone depletion may significantly shorten this season. Although the apparent impact would be lessened by the decrease in UV with depth, irreversible detrimental effects would probably occur before reported survival thresholds were exceeded.Contribution No. 1107 from the Department of Oceanography, University of Washington, Seattle, WA 98195, USA     

Vertical movement of mud crab megalopae (Scylla serrata) in response to light: Doing it differently down under

Selective tidal-streaming is a model frequently used to explain how planktonic larvae invade estuaries. The ability of larvae to move vertically in the water column to selectively ride favourable currents and maintain ground gained is critical to this process. The mud crab (Scylla serrata) is a widely distributed, commercially and recreationally important portunid crab but little is known about its estuarine recruitment mechanisms or the vertical migration behaviour of its megalopae. In studies of the blue crab (Callinectes sapidus), important factors identified in the recruitment mechanism include altered vertical swimming behaviours in estuarine and offshore water and an endogenous circadian rhythm. Using laboratory experiments we examined the vertical displacement response of mud crab megalopae to illumination in estuarine and offshore water during the day and the night. Mud crab megalopae released into 1 m high towers swam higher when illuminated than when in darkness. This behaviour was repeated during the day and the night and in offshore and estuarine water. Given the apparent indifference to water type and the fact that mud crab megalopae are rarely caught in estuaries, we propose the model that these crabs do not invade estuaries as megalopae, but settle and metamorphose into small crabs on the coastal shelf before moving along the sea bed into estuarine habitats.     

Trophodynamics of the plankton community at Dogger Bank: predatory impact by larval fish

The trophodynamics of a coastal plankton community were studied,focusing on fish larvae and their copepod prey. The major objectiveswere to describe distributional overlap and evaluate the predatoryimpact by larval fish. The study was carried out across DoggerBank in the North Sea, August-September 1991. Sampling transectscrossed tidal fronts off the Bank and plankton at all trophiclevels showed peak abundance within frontal zones. Also Verticallythere was a significant overlap in distributional patterns ofthe plankton. Seven species of fish larvae were abundant, ofthese sprat (Sprattus sprattus) dominated. The abundance ofone group of fish larvae peaked in the shallow water close tothe Bank, whereas other species, including sprat, were foundin deeper water. Prey preference and predation pressure of fishlarvae were assessed using information on prey sizes and growthrates of larvae and the copepod prey. We estimated larval removalof preferred prey sizes to 3–4% day^–1, counterbalancedby a 3–7% day^–1' replenishment from copepod productionand growth. Additional predation pressure on copepods by aninvertebrate predator was estimated to 1–3%day^–1.In conclusion, the dynamics of fish larvae and other zooplankterswere closely linked. At peak abundances of fish larvae (>35mg dry weight m^–2), the accumulated predation on specificsize ranges of copepods, exerted by larvae and other predators,could exceed the ability of copepod replenishment and intra-/interspecificcompetition among predators might take place.     

Avoidance of hydrodynamically mixed environments by <Emphasis Type="Italic">Mnemiopsis leidyi</Emphasis> (Ctenophora: Lobata) in open-sea populations from Patagonia,Argentina

Biomass distributions of the ctenophore, Mnemiopsis leidyi, were collected during a week-long survey program in a tidal front along the Patagonian shelf in December 1989. Average ctenophore biomass concentrations varied significantly along a north–south gradient and in stratified compared to unstratified waters. The relative vertical distribution of M. leidyi biomass appeared to be constrained by surface levels of vertical shear. Vertical distributions of ctenophore biomass were highly variable at low levels of vertical shear (<4 s⁻¹) at the surface, but at higher levels of surface vertical shear ctenophores occurred deeper in the water column where shear levels were lower. These results indicate that physical conditions are important factors influencing the distribution of M. leidyi along the Patagonian shelf during summer months.     

Mosquito oviposition and larvae development in response to kairomones originated by different fish

Predator‐released kairomones indicating the presence of predatory fish are known to alter the behaviour and life‐history traits of several aquatic herbivores. Culex mosquitoes (Diptera: Culicidae) respond to such cues by altering oviposition habitat selection and larval development time. These responses differ among fish species indicating composition differences among fish‐released cues, but the recognition pattern is not clear. This study tested the dependence of fish recognition to co‐evolution and the level of threat to larvae. We compared Culex responses to three fish, the invasive larvivorous Gambusia affinis (Cyprinodontiformes: Poeciliidae), the native larvivorous Aphanius mento (Cyprinodontiformes: Cyprinodontidae) and the native algivorous Garra rufa (Cypriniformes: Cyprinidae). The effects of cues released by the three fish on Culex oviposition and life‐history shifts were compared in field mesocosms and lab tests. Our results showed that ovipositing females were significantly repelled only by cues originated from G. affinis, while developing larvae response was a more general one. Our results support the idea that fish‐released kairomones differ in composition or quantities among different species.     

Predator regime influences innate anti-predator behaviour in the freshwater gastropod Lymnaea stagnalis

1. Predation incurs high fitness costs in aquatic organisms either through direct consumption or through avoidance responses that reduce time for activities such as feeding and reproduction. Hence, avoidance responses of aquatic organisms should vary to match closely the predation threat in their environment.
2. The freshwater gastropod Lymnaea stagnalis occurs in a variety of environments which vary in the presence or absence of predatory fish. We used naïve snails reared from six populations of this species experiencing different predator regimes (three co-occurring with molluscivorous fish and three without) to assess whether populations differed in the type and degree of their avoidance behaviours. Innate behavioural responses to four treatments (control, conspecific alarm cues, fish kairomones and fish kairomones paired with alarm cue) were compared in laboratory trials.
3. The primary anti-predator behaviour of L. stagnalis in response to fish kairomones was to crawl out of the water rather than seek refuge under water. This response was strongest when fish kairomones were paired with alarm cues, and varied depending on population origin; snails reared from populations co-occurring with predatory fish showed a stronger response than those raised from populations not experiencing such predators. In addition, populations co-occurring with predatory fish responded to the fish kairomones presented alone.
4. Our findings suggest that the degree of innate anti-predator behaviour shown by L. stagnalis , in terms of both the level of risk to which it responds and the degree of response, varies depending on the predator regime experienced by field populations. Together with previous work on cue association, this demonstrates that this gastropod is able to match its avoidance behaviour very closely to short and long term predation threats within its habitat.     

Chemical composition and oxygen consumption rates of the ctenophore Bolinopsis infundibulum from the Gulf of Maine

Quantitative determinations of chemical composition and oxygenconsumption rates were made for a deep-living population ofthe lobate ctenophore Bolinopsis infundibulum. Animals werecollected in the Gulf of Maine with the submersible ‘Johnson-Sea-Link’during September 1989 at depths ranging from 120 to 240 m. Carbonand nitrogen contents were similar to values reported for epipelagicctenophores. Lipid and protein levels were lower than valuestypical of epipelagic ctenophores, but higher than those ofmesopelagic species. Carbohydrate was nearly an order of magnitudehigher than previously recorded for B.infundibulum. Oxygen consumptionrates ranged from 0.004 to 0.235 µl O₂ mg^–1 dryweight h^– at temperatures ranging from 5 to 7°C. Carbon-specificmetabolic rates ranged from 0.21 to 12.73 µl O₂ mg^–1C h^–1. Energy expenditures estimated from respirationdata (     

Exploring the potential of small RNA subunit and ITS sequences for resolving phylogenetic relationships within the phylum Ctenophora

Ctenophores are a phylum of non-bilaterian marine (mostly planktonic) animals, characterised by several unique synapomorphies (e.g., comb rows, apical organ). Relationships between and within the nine recognised ctenophore orders are far from understood, notably due to a paucity of phylogenetically informative anatomical characters. Previous attempts to address ctenophore phylogeny using molecular data (18S rRNA) led to poorly resolved trees but demonstrated the paraphyly of the order Cydippida. Here we compiled an updated 18S rRNA data set, notably including a few newly sequenced species representing previously unsampled families (Lampeidae, Euryhamphaeidae), and we constructed an additional more rapidly evolving ITS1 + 5.8S rRNA + ITS2 alignment. These data sets were analysed separately and in combination under a probabilistic framework, using different methods (maximum likelihood, Bayesian inference) and models (e.g., doublet model to accommodate secondary structure; data partitioning). An important lesson from our exploration of these datasets is that the fast-evolving internal transcribed spacer (ITS) regions are useful markers for reconstructing high-level relationships within ctenophores. Our results confirm the paraphyly of the order Cydippida (and thus a “cydippid-like” ctenophore common ancestor) and suggest that the family Mertensiidae could be the sister group of all other ctenophores. The family Lampeidae (also part of the former “Cydippida”) is probably the sister group of the order Platyctenida (benthic ctenophores). The order Beroida might not be monophyletic, due to the position of Beroe abyssicola outside of a clade grouping the other Beroe species and members of the “Cydippida” family Haeckeliidae. Many relationships (e.g. between Pleurobrachiidae, Beroida, Cestida, Lobata, Thalassocalycida) remain unresolved. Future progress in understanding ctenophore phylogeny will come from the use of additional rapidly evolving markers and improvement of taxonomic sampling.