Shifting abundance of the ctenophore Mnemiopsis leidyi and the implications for larval bivalve mortality

Along the mid-Atlantic coast of the US, the ctenophore Mnemiopsis leidyi (Agassiz) appears to be increasing in abundance and undergoing shifts in its historical seasonal distribution. We provide new data on shifting ctenophore abundance in Long Island estuaries and its implications for top-down control of the plankton community. Peak mean biovolume estimates of M. leidyi in Long Island estuaries in 2006 revealed ctenophore abundance values that were a factor of two to five times greater than previous studies conducted two decades ago. Furthermore, peak M. leidyi densities in 2006 occurred 2–3 months earlier than previously documented, suggesting a shift in the seasonal maxima of M. leidyi. Application of daily ingestion rates to zooplankton abundance indicates that, at its highest densities M. leidyi can remove an overall average of 20–89% per day of bivalve veligers and other zooplankton taxa, including adult copepods, nauplii, and tintinnids. Increasing ctenophore abundance, especially during a period when they were not historically abundant (i.e., June) may have significant consequences for species which spawn at this time. For example, current populations of M. leidyi represent a major source of larval mortality for bivalves which may inhibit recovery of shellfish populations and reinforce their low abundance state in Long Island estuaries.     

Feeding behavior of the ctenophore <Emphasis Type="Italic">Thalassocalyce inconstans</Emphasis>: revision of anatomy of the order Thalassocalycida

Behavioral observations using a remotely operated vehicle (ROV) in the Gulf of California in March, 2003, provided insights into the vertical distribution, feeding and anatomy of the rare and delicate ctenophore Thalassocalyce inconstans. Additional archived ROV video records from the Monterey Bay Aquarium Research Institute of 288 sightings of T. inconstans and 2,437 individual observations of euphausiids in the Gulf of California and Monterey Canyon between 1989 and 2005 were examined to determine ctenophore and euphausiid prey depth distributions with respect to temperature and dissolved oxygen concentration [dO]. In the Gulf of California most ctenophores (96.9%) were above 350 m, the top of the oxygen minimum layer. In Monterey Canyon the ctenophores were more widely distributed throughout the water column, including the hypoxic zone, to depths as great as 3,500 m. Computer-aided behavioral analysis of two video records of the capture of euphausiids by T. inconstans showed that the ctenophore contracted its bell almost instantly (0.5 s), transforming its flattened, hemispherical resting shape into a closed bi-lobed globe in which seawater and prey were engulfed. Euphausiids entrapped within the globe displayed a previously undescribed escape response for krill (‘probing behavior’), in which they hovered and gently probed the inner surfaces of the globe with antennae without stimulating further contraction by the ctenophore. Such rapid bell contraction could be effected only by a peripheral sphincter muscle even though the presence of circumferential ring musculature was unknown for the Phylum Ctenophora. Thereafter, several live T. inconstans were collected by hand off Barbados and microscopic observations confirmed that assumption.     

Distribution and abundance of ctenophores and their zooplankton food in the Black Sea. II. Mnemiopsis leidyi

The distribution of Mnemiopsis leidyi Agassiz, 1865 in the Black Sea was determined using plankton samples collected above the anoxic zone (maximum depth 200 m) in the summer, winter, and spring from 1991 to 1995. Distribution was patchy. Average biomasses of 15 to 500 g m⁻² were measured, and abundances varied from 10 to 180 ind m⁻². Biomass and abundance peaked in winter, and there was a secondary peak in the summer. The distribution of M. leidyi was correlated with hydrographic features in the Black Sea with higher concentrations in anticyclonic gyres. The centers of the two main cyclonic gyres generally had a low biomass of M. leidyi. From July 1992 to March 1995, the populations were largely offshore. M. leidyi were confined to the upper part of the mixed layer both day and night. Some individuals displayed a negative taxis to daylight and were concentrated below the thermocline at night. Smaller M. leidyi (1.5 to 2 cm) were present in the winter, and individuals reached maximum size in the summer. Although reproduction was continuous throughout the year, there were two distinct peaks: the larger peak in the summer and the smaller peak in the winter. Microscopic analysis of stomach contents showed that copepods and molluscs form their main diet. Received: 1 November 1997 / Accepted: 30 August 1999     

Distribution of gelatinous macrozooplankton in the southern Black Sea during 1996–1999

In this study, spatial and temporal distributions of three gelatinous macrozooplankton, the scyphozoan Aurelia aurita and the ctenophores Pleurobrachia pileus and Mnemiopsis leidyi, were evaluated using the data obtained from seven southern Black Sea cruises carried out from 1996 to 1999. A comparison of nets used for the sampling of gelatinous macrozooplankton illustrated the superiority of the Hensen net with a larger mesh size (300 µm) over the Nansen net (112 µm). P. pileus was mainly concentrated below the mixed layer, while A. aurita and M. leidyi were generally confined to surface waters. Horizontally, P. pileus was associated with deep, offshore waters. P. pileus also displayed the least interannual variation of the three species. Biomass of A. aurita and M. leidyi varied substantially seasonally and inter-annually. In September 1999, the average biomass of M. leidyi was at its lowest level (12 g wet weight m^-2) since its explosive development at the end of the 1980s. This was associated with the appearance of the new predatory ctenophore Beroe ovata, which appeared in the Black Sea at the end of the 1990s.     

Distribution of the alien ctenophore <Emphasis Type="Italic">Mnemiopsis</Emphasis><Emphasis Type="Italic">leidyi</Emphasis> in the Caspian Sea in August 2001

In this study, spatial and vertical distributions of the invasive ctenophore Mnemiopsis leidyi in the Caspian Sea were evaluated by using data collected at 41 stations during the August 2001 cruise. A comparison of data from different depths revealed that M. leidyi were generally confined to surface waters. The maximum size of the ctenophore was only 41-45 mm, and the bulk of individuals (85.5%) were <10 mm in length. The average and maximum biomasses of M. leidyi were calculated as 120 and 351 g wet weight m^-2, respectively. Whilst highest biomasses were observed in the western and central Middle Caspian Sea, hot spot areas of reproduction were present along the coasts of the western Caspian Sea, with abundance values of up to 2285 ind. m^-2. The impact of such high densities of M. leidyi is expected to be significant for the pelagic ecosystem of the Caspian Sea.     

Population development of the invader ctenophore Mnemiopsis leidyi, in the Black Sea and in other seas of the Mediterranean basin

In the last two decades of the twentieth century, the ctenophore Mnemiopsis leidyi (A. Agassiz) has invaded the Black, Azov, Marmara and Aegean Seas, and, recently, the Caspian Sea. Here, we compare its spatial and temporal distribution, seasonal dynamics and the time and duration of reproduction. We also discuss factors that control its abundance throughout its invasive range and its effect on ecosystems. Observations are based on the long-term field data collected by three research institutes. An analysis of the effects of temperature, salinity, prey (zoo- and ichthyoplankton) availability and predation (by ctenophores of the genus Beroe) on M. leidyi population size, and the effects of M. leidyi on zoo- and ichthyoplankton, and on fish populations in the Black and Azov Seas is also provided. With the Black Sea current, M. leidyi spreads to the upper layers of the Sea of Marmara, where it now occurs around the year. At regular intervals, the Black Sea current also takes it to the northern Aegean Sea. In contrast, it has to re-invade the Sea of Azov every spring or summer, dying out during winter when the temperature drops below 4°C. The warm summer and mild winter temperatures, relatively low salinity and abundance of prey in the Black Sea are close to optimal for M. leidyi, while they are suboptimal in the northern Aegean Sea, where salinity and temperature are often too high. In the Black Sea the absence of gelatinous and other predators led to an enormous ctenophore abundance for a decade, but with the appearance of Beroe ovata in 1999, M. leidyi abundance greatly decreased. Analysis of seasonal dynamics of M. leidyi in the Black Sea and in other seas of the Mediterranean basin indicates similarities in the timing of maximum abundance and biomass, in spite of some differences in the initiation and duration of reproduction. A peak biomass and density occurred in 1989 in the Black and Azov Seas and in 1990 in the other seas. The M. leidyi invasion negatively affected the ecosystems of the Black Sea and the Sea of Azov. The zooplankton, ichthyoplankton and zooplanktivorous fish stocks all underwent profound changes. Similar effects, but less pronounced, were recorded in the Sea of Marmara. Effects on Mediterranean food chains have, so far, remained insignificant. Salinity is probably supraoptimal here, and several predators prevent M. leidyi from reaching outbreak levels.     

Ingestion of a medusa (Aegina citrea) by the nematocyst-containing ctenophore Haeckelia rubra (formerly Euchlora rubra): phylogenetic implications

The rare ctenophore Haeckelia rubra (formerly Euchlora rubra) has long been known to have nematocysts rather than colloblasts in its tentacles. Five specimens were collected in the San Juan Archipelago, Washington State, USA in 1980 and 1981, and their feeding behavior was observed in the laboratory. We found that H. rubra readily eats the tentacles of a medusa, Aegina citrea, whose nematocysts (apotrichous isorhizas) are nearly identical in morphology to the nematocysts of the ctenophore. When H. rubra was offered 16 other species of hydromedusae and 1 siphonophore in the laboratory, the ctenophores showed little or no tendency to ingest these potential prey items. In addition to its routinely positive response to A. citrea, the ctenophore could be induced by manipulation and starvation to accept and ingest bits of the bodies of 4 additional species of hydromedusae and 1 siphonophore. These results, combined with the histological and rearing experiments of other investigators, leave little doubt that the nematocysts in H. rubra are not endogenous, but are kleptocnidae similar to those nematocysts retained and subsequently used by some species of nudibranchs that feed on Cnidaria. A close phylogenetic link between the Cnidaria and the Ctenophora is most unlikely.     

Ammonium excretion by gelationous zooplankton and their contribution to the ammonium requirements of microplankton in Chesapeake Bay

Ammonium excretion rates of recently collected specimens of gelatinous zooplankton, the scyphomedusan Chrysaora quinquecirrha DeSor and the etenophore Mnemiopsis leidyi A. Agassiz, were correlated with body mass and water temperature in measurements made from April to October 1989 and 1990. Rates ranged between 3.5 and 5.0 g atoms NH ₄ ⁺ -N (g dry wt)^-1h^-1 for C. quinquecirrha and 3.0 to 4.9 g atoms NH ₄ ⁺ -N (g dry wt)^-1h^-1 for M. leidyi. Excretion rate equations and in situ data on the size distributions and biomasses of gelatinous zooplankters and water temperature were used to estimate the contribution of ammonium by medusae and ctenophores to mesohaline Chesapeake Bay waters on several dates during April to October 1989 and 1990. We then compared the estimated contributions to direct measurements of ¹⁵NH ₄ ⁺ uptake by microplankton. The maximum estimated regeneration by gelatinous zooplankton was 5.8 g atoms NH ₄ ⁺ -N m^-3h^-1 at night in August 1990, when medusae biomass was greatest. This represents about 4% of the ammonium required by the microplankton. During the daytime on all dates, less than 1% of the ammonium required by microplanktion was supplied by gelatinous zooplankton. Therefore, gelatinous zooplankton appear to play a minor role in the ammonium cycle of Chesapeake Bay.     

A three-trophic level estuarine model: Synergism of two mechanistic simulations

Few numerical simulations have attempted to include a high degree of biological detail for several trophic levels. Typically, in planktonic ecosystem models, if the dynamics of nutrients, phytoplankton and herbivorous zooplankton are formulated with ecological complexity, then carnivores are ignored, forced or modeled in an extremely simplified manner. Extensive mechanistic detail for important carnivores is difficult to represent because reliable and relevant ecological data are rarely available for appropriate species and local populations. Further, the wide temporal and spatial differences between life histories of lower plankton and carnivores may be technically difficult to model.In Narragansett Bay, Rhode Island, the ctenophore Mnemiopsis leidyi is an important carnivore to which these objections do not apply. A detailed carbon-based simulation model of this population of ctenophores was developed independently from an ecosystem model of Narragansett Bay which included detailed interactions between phytoplankton, primarily herbivorous zooplankton and nutrients. The interfacing of these two models without changing any of the formulations or values of the coefficients provided a test of the commonly used practice of forcing certain components. Both models were originally constructed with the biomass of a critical compartment forced according to observed data; in the plankton model, ctenophores were forced, and in the ctenophore model, zooplankton were forced.Predicted biomasses for zooplankton and ctenophores in the combined model were similar to the results of the two parent models, but improved relative to the actual field observations. From the findings it appears that the strategy of forcing is valid provided the forced patterns are appropriate and reasonable.     

Recent distribution and size structure of gelatinous organisms in the southern Black Sea and their interactions with fish catches

The spatial distributions of gelatinous organism were studied during three cruises in 2006–2007. These were the first such studies conducted in the southern Black Sea for last decade. Additionally, the different methods of estimating gelatinous organism biomass presently in use were compared and recent situation of pelagic fishery interacting with zooplankton in the Black Sea were overviewed. Biomasses and abundances of both invading Mnemiopsis leidyi, and the resident Aurelia aurita and Pleurobrachia pileus were remarkably reduced (total gelatinous biomass: 417–537 g m⁻² in May, June, and 150 g m⁻² in October) as compared with those before the 1997 with invasion Beroe ovata in the southern Black Sea. Specimens of B. ovata were detected only in October when individuals with lengths of 10–50 mm comprised 93% of its population. Length–weight based biomasses were significantly variable for all species depending on the size structure in time. Larger sized individuals (>30 mm) of M. leidyi appeared as compared with those before year 1997. The ecosystem of the Black Sea has been suppressed both bottom–up through decreased eutrophication and top–down through reduced grazing on mesozooplankton by M. leidyi that are in turn controlled by grazing by B. ovata. The catch of anchovy was abruptly reduced to 120,000 tons due presumably to the considerably increased catch of its predator, the bonito (64,000 tons) in the year 2005.     

Vulnerability of the copepod Acartia tonsa to predation by the scyphomedusa Chrysaora quinquecirrha : effect of prey size and behavior

Although scyphomedusae have received increased attention in recent years as important predators in coastal and estuarine environments, the factors affecting zooplankton prey vulnerability to these jellyfish remain poorly understood. Current models predicting feeding patterns of cruising entangling predators, such as Chrysaora quinquecirrha (Desor, 1948), fail to account for the selection of fast-escaping prey such as copepods. Nevertheless, our analysis of gastric contents of field-collected medusae showed that this scyphomedusa fed selectively on the calanoid copepod Acartia tonsa (Dana, 1846) and preferentially ingested adult over copepodite stages. We measured feeding rates in a planktonkreisel while simultaneously videotaping predator–prey interactions. C. quinquecirrha consumed adult A. tonsa ten times faster than copepodites. Differences in prey behavior, in the form of predator–prey encounter rates or post-encounter escape responses, could not account for the elevated feeding rates on adults. Prey size, however, had a dramatic impact on the vulnerability of copepods. In experiments using heat-killed prey, feeding rates on adults (1.5 times longer than copepodites) were 11 times higher than on copepodites. In comparison, medusae ingested heat-killed prey at only two to three times the rate of live prey. These results suggest that during scyphomedusan–copepod interactions, prey escape ability is important, but ultimately small size is a more effective refuge from predation. Received: 26 September 1997 / Accepted: 20 May 1998     

Grazing of phytoplankton by copepods in eastern Antarctic coastal waters

Chlorophyll a, primary productivity and grazing by copepods on phytoplankton were measured in the upper water column during the summer of 1994/1995 at a coastal site near Davis Station, East Antarctica. Chlorophyll a was at a maximum in mid-December, then dropped markedly as the coastal fast ice melted and broke‐out. Phytoplankton biomass increased again from mid‐ to late‐February. Copepods accounted for at least 65% of zooplankton biomass in the water column before sea ice break‐out, whereas larval polychaetes and ctenophores dominated after ice break‐out. Oncaeacurvata was the numerically dominant species throughout the study. The highest grazing rate (8.7 mg C␣m⁻³␣d⁻¹) was recorded on 21 December when O.␣curvata accounted for 64% of the total. Grazing had decreased markedly by 28 December (0.9 mg C m⁻³ d⁻¹); again O. curvata accounted for over 50% of the total ingested. Copepod grazing increased after ice break-out until the last experiment on 20 February (⋍5 mg C␣m⁻³␣d⁻¹). The main species responsible for grazing during this period were O. curvata, Oithonasimilis, Calanoidesacutus and unidentified copepod nauplii. It was estimated that copepods removed between 1 and 5% of primary productivity. Received: 11 October 1996 / Accepted: 22 October 1996     

Prey, feeding rates, and asexual reproduction rates of the introduced oligohaline hydrozoan Moerisia lyonsi

Moerisia lyonsi Boulenger (Hydrozoa) medusae and benthic polyps were found at 0 to 5‰ salinity in the Choptank River subestuary of Chesapeake Bay, USA. This species was introduced to the bay at least 30 years before 1996. Medusae and polyps of M. lyonsi are very small and inconspicuous, and may occur widely, but unnoticed, in oligohaline waters of the Chesapeake Bay system and in other estuaries. Medusae consumed copepod nauplii and adults, but not barnacle nauplii, polychaete and ctenophore larvae or tintinnids, in laboratory experiments. Predation rates on copepods by medusae increased with increasing medusa diameter and prey densities. Feeding rates on copepod nauplii were higher than on adults and showed no saturation over the range of prey densities tested (1 to 64 prey l⁻¹). By contrast, predation on copepod adults was maximum (1 copepod medusa⁻¹ h⁻¹) at 32 and 64 copepods l⁻¹. Unexpectedly, M. lyonsi colonized mesocosms at the Horn Point Laboratory during the spring and summer in 4 years (1994 to 1997), and reached extremely high densities (up to 13.6 medusae l⁻¹). Densities of copepod adults and nauplii were low when medusa densities were high, and estimated predation effects suggested that M. lyonsi predation limited copepod populations in the mesocosms. Polyps of M. lyonsi asexually produced both polyp buds and medusae. Rates of asexual reproduction increased with increasing prey availability, from an average total during a 38 d experiment of 9.5 buds polyp⁻¹ when each polyp was fed 1 copepod d⁻¹, to an average total of 146.7 buds polyp⁻¹ when fed 8 copepods d⁻¹. The maximum daily production measured was 8 polyp buds and 22 medusae polyp⁻¹. The colonizing potential of this hydrozoan is great, given the high rates of asexual reproduction, fairly wide salinity tolerance, and existence of a cyst stage. Received: 29 October 1998 / Accepted: 3 March 1999     

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

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

Swimming and feeding by the scyphomedusa Chrysaora quinquecirrha

The semaeostome scyphomedusa, Chrysaora quinquecirrha (Desor, 1848), is an abundant and important planktonic predator in estuaries and coastal waters of the eastern USA during the summer. We videotaped free-swimming medusae in the laboratory and in the field in order to determine the relationship between swimming motions and prey encounter with capture surfaces. Medusae were collected from the Choptank River (Chesapeake Bay) in September 1992 and in the Niantic River, Connecticut, USA in July 1994. We used newly hatched Artemia sp. nauplii and fluorescein dye to trace water motions around swimming medusae. Swimming results in a pulsed series of toroids which travel along the medusan oral arms and tentacles. Prey are entrained in this flow and the location of naupliar encounter was influenced by the phase of the pulsation cycle during which entrainment occurred. Flow-field velocities, measured by tracking particles adjacent to the bell margin during contraction, increased with bell diameter. Received: 29 March 1997 / Accepted: 11 April 1997     

Population biology and antipredator defenses of the shallow-water Antarctic nudibranch Tritoniella belli

The circumpolar nudibranch Tritoniella belli Eliot occurs in abundance in shallow-water benthic communities of McMurdo Sound, Antarctica. Density estimates based on belt transects averaged collectively 0.46 individuals m⁻² at three study sites between depths of 6 and 30 m in November 1996. At two of the sites, population densities increased linearly between 18 and 30 m depth (up to 0.7 and 1.15 individuals m⁻² at 30 m depth). Individuals at all sites were rare or absent at depths shallower than 12 m. Size frequencies of individuals at the sites were similar, and a pooled analysis revealed a unimodal distribution skewed highly towards juvenile size classes. This suggests both recent recruitment and constant rates of mortality across size classes. The relationship between foot length and wet weight best fits an exponential growth equation, indicative of an allometric growth pattern. Distribution of T. belli in the field suggests that it is a habitat and diet generalist. Potential invertebrate predators include sea anemones and seastars, both of which co-occur in abundance in McMurdo Sound. Laboratory experiments indicate that the sea anemone Isotealia antarctica can capture and ingest T. belli. However, 70% of T. belli that are captured escape from the tentacles or, following ingestion, are rejected from the gastrovascular cavity. The seastars Odontaster validus, Perknaster fuscus, and Acodontaster conspicuus, avoid contact with T. belli, but if forced into contact with mantle tissues, retract their tube-feet. Mucus secreted from the mantle tissues, coated on to the tips of glass rods, and presented to seastar tube-feet, causes significantly longer tube-foot retraction times than control rods. Moreover, pieces of freeze-dried krill coated with mantle mucus are consumed significantly less often than untreated control pieces of krill by a benthic scavenging fish (Pseudotrematomas bernacchi). Employing seastar tube-foot retractions as a bioassay, we found the bioactive compound(s) are soluble in ethyl acetate, indicating they are lipophilic or moderately hydrophilic in nature. Chemical defenses in the mucus of T. belli probably contribute to its high abundance in Antarctic benthic communities. Received: 6 October 1997 / Accepted: 24 March 1998     

Nucleic acid and protein levels in strobilating polyps of Chrysaora quinquecirrha and Aurelia aurita

Levels of DNA, RNA, and protein were measured in scyphistomae of the scyphozoans Aurelia aurita and Chrysaora quinquecirrha during strobilation. In synchronously developing populations of A. aurita, the amount of DNA per polyp increases 41 to 260% during strobilation, whereas RNA and protein remain relatively constant. The RNA/DNA ratio drops by about 50% during strobilation in both C. quinquecirrha and A. aurita, and protein/DNA decreases by 50% in A. aurita. Specific RNA reserves are probably not accumulated in advance of strobilation, since strobilation is blocked immediately in both species by 1 to 3 g/ml actinomycin D. The size and DNA content of polyps are usually increased by prolonged incubation at temperatures of 12° to 15°C, which facilitates strobilation; however, the RNA/DNA and protein/DNA ratios of such polyps are not significantly increased by the cold conditioning.Contribution No. 472 from the Virginia Institute of Marine Science.     

Foraging ecology of leatherback sea turtles in the Western North Atlantic determined through multi-tissue stable isotope analyses

Leatherback turtles, Dermochelys coriacea, are highly migratory, spending most of their lives submerged or offshore where their feeding habits are difficult to observe. In order to elucidate the foraging ecology of leatherbacks off Massachusetts, USA, stable isotope analyses were performed on leatherback tissues and prey collected from 2005 to 2009. Stable isotope ratios of nitrogen and carbon were determined in whole blood, red blood cells, blood plasma, muscle, liver, and skin from adult male, female, and subadult leatherbacks. Isotopic values were analyzed by body size (curved carapace length) and grouped by sex, and groups were tested for dietary differences. Gelatinous zooplankton samples were collected from leatherback foraging grounds using surface dip nets and stratified net tows, and prey contribution to leatherback diet was estimated using a two-isotope Bayesian mixing model. Skin and whole blood δ¹³C values and red blood cell δ¹⁵N values were correlated with body size, while δ¹³C values of red blood cells, whole blood, and blood plasma differed by sex. Mixing model results suggest that leatherbacks foraging off Massachusetts primarily consume the scyphozoan jellyfishes, Cyanea capillata and Chrysaora quinquecirrha, and ctenophores, while a smaller proportion of their diet comes from holoplanktonic salps and sea butterflies (Cymbuliidae). Our results are consistent with historical observations of leatherback turtles feeding on scyphozoan prey in this region and offer new insight into size- and sex-related differences in leatherback diet.     

Effects of contrasting modes of larval development on the genetic structures of populations of three species of prosobranch gastropods

 In south-eastern Australia, the prosobranch gastropods Morula marginalba (Blainville), Cominella lineolata (Lamarck) and Bedeva hanleyi (Angas) have similar fine-scale distributions, but appear to possess very different dispersal capabilities due to contrasting modes of larval development. M.marginalba produce planktonic larvae, whereas C. lineolata and B. hanleyi undergo direct development in benthic egg capsules and emerge as crawling juveniles. To test for possible effects of contrasting life histories on levels of genetic variation within and among populations, a survey was conducted of allozyme variation at six polymorphic loci in 8 to 9 local populations of each species. Collections of snails were made between June 1992 and November 1993. Sampling ranges spanned between 162 and 180 km of coast. Regardless of larval type, proportions of single-locus genotypes in each collection were consistent with the recruitment of offspring which had been generated through random mating. However, genotypic diversity was lower in those species that undergo direct development. Loci surveyed in C. lineolata and B. hanleyi were polymorphic (i.e. frequency of most common allele <95%) in fewer populations than those examined for M.␣marginalba (P <0.001) and, where polymorphisms occurred, also possessed significantly fewer alleles (P <0.001). Consequently, average levels of expected heterozygosity were greater in populations of M. marginalba than in those of either of the other species (P <0.001). Genetic variation among populations, expressed as the standardised variance in allele frequencies (F _ST ), was inversely related to expected larval dispersal capability. The nine collections of M. marginalba showed little overall differentiation (F _ST  = 0.017; P <0.001), reflecting the ability of planktonic larvae to interconnect local populations, and so limit divergence due to drift and natural selection. In contrast, there were high levels of allelic heterogeneity among the nine collections of C. lineolata (F _ST  = 0.523; P <0.001) and eight collections of B. hanleyi (F _ST  = 0.140; P <0.001). These data imply that for species which undergo direct development, local populations are effectively closed and evolve largely independent of one another. Received: 3 May 1996 / Accepted: 12 July 1996     

Metabolism of epipelagic tropical ctenophores

Measurements of respiration and excretion at 25°C were made for five species of ctenophores collected during five cruises to the Bahamas (1982–1984). The mean element-specific respiration and ammonium excretion rates of freshly collected specimens of all species ranged from 4 to 16% d^-1, the mean atomic O:N ratios were 10 to 16, and ammonium averaged 60 to 90% of the total dissolved nitrogen excreted. For adult ctenophores, the carbon content ranged from 0.6% carbon (as percent of dry weight) for Bolinopsis vitrea to 3.7% carbon for Beroë ovata. There was a marked increase in the organic content (% carbon of dry weight) of small Bolinopsis vitrea with tentacles compared to fully lobate adults. B. vitrea had increasingly higher metabolic rates when held at food concentrations up to 100 copepods 1^-1 (about 250 g C 1^-1). The overall range between starved and well-fed B. vitrea was about two times for respiration and a factor of three for ammonium excretion. B. vitrea decreased from well-fed to a starved metabolic rate in about a day after removal from food. The metabolic rate of Eurhamphaea vexilligera was not measurably affected by short-term starvation or feeding (maximum 25 copepods 1^-1). In feeding experiments, E. vexilligera of 20 to 56 mm length fed at rates equivalent to clearance rates of 250 to 1 800 ml h^-1.