Escape of the ctenophore Mnemiopsis leidyi from the scyphomedusa predator Chrysaora quinquecirrha

The ctenophore Mnemiopsis leidyi A. Agassiz, 1865 is known to be eaten by the scyphomedusan Chrysaora quinquecirrha (Desor, 1948), which can control populations of ctenophores in the tributaries of Chesapeake Bay. In the summer of 1995, we videotaped interactions in large aquaria in order to determine whether M. leidyi was always captured after contact with medusae. Surprisingly, M. leidyi escaped in 97.2% of 143 contacts. The ctenophores increased swimming speed by an average of 300% immediately after contact with tentacles and 600% by mid-escape. When caught in the tentacles of C. quinquecirrha, the ctenophores frequently lost a portion of their body, which allowed them to escape. Lost parts regenerated within a few days. The striking ability of M. leidyi to escape from C. quinquecirrha may be critically important in maintaining ctenophore populations in situ. Received: 14 November 1996 / Accepted: 4 December 1996     

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.     

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.     

A comparative study of zooplankton dynamics in two subtropical temporarily open/closed estuaries, South Africa

This study aims at comparing the zooplankton abundance/biomass of two South African estuaries affected to a different degree by sewage pollution. In particular the hypothesis that microzooplankton may increase relative to mesozooplankton, as eutrophication increases, is tested for the first time in African estuaries. The survey was carried out during a whole-year cycle, including both open and closed phases. Results show that zooplankton abundance and biomass are higher in the Mdloti than in the Mhlanga, during both the closed (U=1, P<0.001, U=16, P<0.001) and the open phase (U=101, P<0.05, U=88, P<0.01). There were no significant differences in abundance/biomass between the different reaches of each estuary, during either their open or closed phase (1-way ANOVA, P>0.05). The different levels of dominance of the calanoid copepod Pseudodiaptomus hessei, which accounted for 53–64 and 86–97% of the total abundance in the Mdloti and the Mhlanga, respectively, suggests a shift in the zooplankton community structure. There was a lower microzooplankton abundance/biomass contribution to the total zooplankton in the Mhlanga (59.6–15.8%), compared to the Mdloti (99.1–96.5%). The highest microzooplankton contribution in the Mdloti was observed during the closed phase, while in the Mhlanga this occurred during the open phase. This suggests that eutrophication in the Mhlanga may impact negatively on the microzooplankton community. On the other hand the less affected Mdloti exhibits an opposite trend with an increased microzooplankton component.     

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 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.     

Deposit-feeding amphipods (Monoporeia affinis) reduce the recruitment of copepod nauplii from benthic resting eggs in the northern Baltic Sea

We experimentally investigated the effect of different densities of the burrowing, deposit-feeding amphipod Monoporeia affinis on the recruitment of zooplankton from benthic resting eggs. Intact sediment cores with in situ density and species composition of zooplankton resting eggs and benthic fauna were collected in the northern Bothnian Sea, part of the Baltic Sea. We removed as many M. affinis as possible from the cores and then added different numbers of M. affinis to the cores to generate a range of densities. The cores were exposed to different densities of M. affinis for either 3 or 40 days, after which the hatched zooplankton was registered. One subset of the cores were initially incubated under low temperature (2–3 °C, to prevent hatching) for 37 days (the resting phase), to allow for effects of M. affinis on unhatched resting eggs. These cores were then incubated under higher temperature (13 °C) for 3 days (the hatching phase), to induce hatching and allow for effects on hatching or hatched specimens. In a second subset of cores with the same time and temperature schedule, the M. affinis density was experimentally reduced at the start of the hatching phase, to evaluate the effect of M. affinis during the hatching phase. To a third subset of cores, we immediately initiated the hatching phase, without an experimental resting phase, to evaluate the effects induced during the resting phase. The most common zooplankton species that hatched was Eurytemora affinis (Copepoda), followed by Bosmina longispina maritima (Cladocera). In all cores that were subjected to a resting phase, the numbers of hatched E. affinis were log-linearly negatively related to density of M. affinis. An increase of M. affinis density from 1,000 to 5,000 individuals m⁻², normal field densities, reduced the hatching by 60–70%. The negative impact was mainly exerted during the hatching phase, suggesting predation on, burial of or physical injury of hatching nauplii or eggs in a late development stage as likely mechanisms. Also, the number of B. longispina maritima that hatched was reduced by M. affinis during the hatching phase, but no clear relation to density of M. affinis could be identified. The results show that M. affinis can reduce recruitment to zooplankton from benthic resting eggs. Such impact by the benthos on resting stages of zooplankton is therefore a potentially significant link between the benthic and pelagic systems. Received: 10 August 2000 / Accepted: 13 November 2000     

Sink or swim? Copepod population maintenance in the Columbia River estuarine turbidity-maxima region

Maintenance of estuarine zooplankton populations in large river-dominated estuaries with short residence times has been an intriguing subject of investigation. During three different hydrological seasons, autumn 1990, summer 1991, and spring 1992, we intensively sampled zooplankton populations in the estuarine turbidity maxima (ETM) region of the Columbia River estuary of Oregon and Washington, USA. One of the principal objectives was to investigate retention mechanisms of the predominant zooplankton species, the harpacticoid copepod Coullana canadensis and the epibenthic calanoid copepod Eurytemora affinis. In the ETM, C. canadensis densities mirrored those of turbidity gradients and were almost always greater at the river bed, while E. affinis densities were greater higher in the water column during the flood and lower in the water column during the ebb. Cross-correlation and time-series analyses determined that C. canadensis densities were highly positively correlated with turbidity and that most of the variability was explained by the lunisolar diurnal (K₁) and principal lunar (M₂) tidal components occurring once every 23.93  h and once every 12.42 h, respectively. This indicates that C. canadensis populations are most probably maintained in the estuary through the same near-bottom circulation features that trap and concentrate particles in the ETM. In contrast, densities of the more motile species E. affinis were highly correlated with negative velocities, or ebb tide, and most of the variability in population densities could be explained by the principal lunar tidal component; therefore, we hypothesize that this species is probably vertically migrating on a tidal cycle into different flow layers to avoid population losses out of the estuary. Received: 7 April 1997 / Accepted: 22 April 1997     

Impact of a burrowing deposit-feeder, Monoporeia affinis, on viable zooplankton resting eggs in the northern Baltic Sea

We investigated the effect of different densities of the burrowing deposit-feeding amphipod Monoporeia affinis on the potential for recruitment of zooplankton from benthic resting eggs. Hatching of resting eggs was induced in the laboratory on sliced and resuspended 1-cm depth-sections of sediment cores, collected at six stations in an archipelago area of the Gulf of Bothnia, Baltic Sea. The uppermost 5 cm of the sediment was studied. The most common species that hatched was Eurytemora affinis (Copepoda). Individuals from another copepod genus, Acartia, hatched in significant numbers only in the cores from two stations with low amphipod abundance. Cores from stations with high amphipod densities showed a deeper distribution of emerging E. affinis nauplii compared with stations with few amphipods; the oxidised sediment layer was also deeper at high M. affinis densities than at low. Total (0 to 5 cm strata pooled) number of hatched E. affinis nauplii was independent of amphipod density. This indicates that the effect of M. affinis on E. affinis eggs involves deeper burial due to bioturbation, rather than predation. Decreased benthic recruitment of zooplankton at localities with high M. affinis density is suggested, since more deeply positioned eggs are less likely to hatch. When hatching was induced in intact, non-sliced cores from one station, the number of E. affinis nauplii that hatched was on average 43% of the number that hatched in the upper centimetre of the sliced cores from the same station. This fraction (43%), if applied to the other stations, implied a potential for benthic recruitment of up to 80 000 ind m⁻² for E. affinis. Due to its high abundance, M. affinis is likely to greatly reduce benthic recruitment of zooplankton in this system. Received: 17 September 1999 / Accepted: 23 February 2000     

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.     

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     

Salps in the Lazarev Sea,Southern Ocean: I. Feeding dynamics

Feeding dynamics of the Antarctic salps Ihlea racovitzai and Salpa thompsoni were studied in the Lazarev Sea in fall 2004, summer 2005–2006 and winter 2006. Pigment concentrations in the guts of both species were positively correlated with ambient surface chlorophyll a (chl a). No evidence was found for salp clogging even at dense surface concentrations of up to 7 μg chl a L⁻¹. However, gut pigment concentrations had a lower range than ambient pigment concentrations, suggesting that salps increased retention times of ingested material in low-food environments. For medium-sized I. racovitzai and S. thompsoni, estimated individual daily rations reached 7–10 and >100% of body carbon in winter and summer, respectively. Daily respiratory needs of I. racovitzai and S. thompsoni accounted for 28 and 22% of daily carbon assimilation based on pigment ingestion rates in winter, and for 2 and 1% in summer, respectively. The grazing impact of the salp populations on the phytoplankton standing stock was negligible during all seasons due to generally low salp densities. Fatty acid trophic biomarkers in the salps suggest high year-round contributions of flagellates and modest contributions of diatoms to the salp’s diet. These markers showed low seasonal variability for I. racovitzai. The more pronounced seasonality of trophic markers in S. thompsoni were likely related to their generally deeper residence depth in winter linked to a seasonal alternation of sexual and asexual generations.     

Seasonal abundance and toxicity of the dinoflagellate Gambierdiscus spp. (Dinophyceae), the causative agent of ciguatera in Tahiti, French Polynesia

The benthic dinoflagellate Gambierdiscus spp. is the primary causative agent of ciguatera fish poisoning in French Polynesia. However, the initiation of ciguatera outbreaks and the factors that control ciguatoxin production in this dinoflagellate are still poorly understood. In this paper, we present some original data concerning the seasonal abundance and toxicity of natural populations of Gambierdiscus spp. based on a long-term survey in a ciguateric site of the island of Tahiti. From February 1993 to December 1997, Gambierdiscus spp. population densities were monitored weekly in the Atimaono barrier reef of Papara in relation to temperature and salinity. Densities peaked at 4992 cells g⁻¹ in October 1994 and constituted ≥1000 cells g⁻¹ on several occasions. A total of 58 blooms were recorded in the area, of which 65% occurred in 1995 and 1996 alone. Our data confirmed the high endemicity of Gambierdiscus spp. in the Papara area. Refined analysis of our raw data by a principal-component analysis of processes (PCAP) revealed a seasonality in cell densities from February 1993 to May 1995. During this period, Gambierdiscus spp. populations tended to reach maximum abundance at the beginning and end of the hot season. In contrast, salinity did not appear to be a determining factor in the seasonal abundance of this dinoflagellate. The noticeable increase in both peak densities and frequency of blooms further noticed in 1995 and 1996 was preceded by unusually high water temperatures in January to April 1994, concomitant with a severe coral-bleaching episode. During the course of the study, a total of 303 × 10⁶ cells obtained from 48 successive blooms was harvested. Toxicity screening revealed that toxin production was maximum from October 1994 through December 1996. No correlation was found between toxicity of these blooms and their biomass nor the seasonal pattern of temperatures. It is suggested that the toxicity of naturally-occurring blooms of Gambierdiscus spp. and, consequently, the severity of ciguatera incidents in a given area, is mainly dependent on the clonal nature of cells which coexist within local populations of this dinoflagellate. Received: 12 October 1998 / Accepted: 15 June 1999     

Estimating abundance of killer whales in the nearshore waters of the Gulf of Alaska and Aleutian Islands using line-transect sampling

Killer whale (Orcinus orca Linnaeus, 1758) abundance in the North Pacific is known only for a few populations for which extensive longitudinal data are available, with little quantitative data from more remote regions. Line-transect ship surveys were conducted in July and August of 2001–2003 in coastal waters of the western Gulf of Alaska and the Aleutian Islands. Conventional and Multiple Covariate Distance Sampling methods were used to estimate the abundance of different killer whale ecotypes, which were distinguished based upon morphological and genetic data. Abundance was calculated separately for two data sets that differed in the method by which killer whale group size data were obtained. Initial group size (IGS) data corresponded to estimates of group size at the time of first sighting, and post-encounter group size (PEGS) corresponded to estimates made after closely approaching sighted groups. ‘Resident’-type (fish-eating) killer whales were more abundant than the ‘transient’-type (mammal-eating). Abundance estimates of resident killer whales (991 [95% CI = 379–2,585] [IGS] and 1,587 [95% CI = 608–4,140] [PEGS]), were at least four times greater than those of the transient killer whales (200 [95% CI = 81–488] [IGS] and 251 [95% CI = 97–644] whales [PEGS]). The IGS estimate of abundance is preferred for resident killer whales because the estimate based on PEGS data may show an upward bias. The PEGS estimate of abundance is likely more accurate for transients. Residents were most abundant near Kodiak Island in the northern Gulf of Alaska, around Umnak and Unalaska Islands in the eastern Aleutians, and in Seguam Pass in the central Aleutians. This ecotype was not observed between 156 and 164°W, south of the Alaska Peninsula. In contrast, transient killer whale sightings were found at higher densities south of the Alaska Peninsula between the Shumagin Islands and the eastern Aleutians. Only two sightings of ‘offshore’-type killer whales were recorded during the surveys, one northeast of Unalaska Island and the other south of Kodiak Island. These are the first estimates of abundance of killer whale ecotypes in the Aleutian Islands and Alaska Peninsula area and provide a baseline for quantifying the role of these top predators in their ecosystem. Electronic Supplementary Material  Supplementary material is available in the online version of this article at and is accessible for authorized users.

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Zooplankton community structure across an eddy-generated upwelling band close to a tropical bay-mangrove ecosystem

Mesoscale eddies in the world’s oceans are ubiquitous and bring about episodic pulses of nutrients into the photic zone. Transient in nature, the role of eddy pumping in coastal enrichment via plankton production, and subsequent organic flux is not yet fully realised. In the context of a cyclonic cold-core eddy that propagates annually under the influence of the East India Coastal Current and enriches coastal waters in the western Bay of Bengal north of 16°N, a detailed study on zooplankton community structure along with phytoplankton composition and associated water quality was undertaken during April–May 2002 coinciding with the spring intermonsoon. Zooplankton samples were collected at 32 hydrographically different (salinity 24.5–35.6 PSU) GPS fixed locations representing bay-mangrove areas and nearshore waters (30 m) close to the River Godavari, which is one of India’s largest estuarine systems. During the study, the bay-coastal waters were typified by elevated nutrient levels (nitrate 10.73–22.04 μM), high salinity (27.98–35.52 PSU), and relatively low temperatures (30.63–31.40°C). Altogether, 95 zooplankton taxa were encountered with copepods forming the predominant population. Agglomerative Hierarchical Cluster Analysis (AHCA) and Non-metric Multidimensional Scaling (NMDS) based on Bray–Curtis similarity (PRIMER) analysis revealed appreciable alterations in zooplankton structure across bay-mangrove locations and coastal waters (Stress 0.11; ANOSIM test Global R: 0.94, P = 0.1%). Similarity Percentage (SIMPER) analysis revealed zooplankton associations through “discriminating species” for each location (Kakinada Bay, Cluster I, 27.9 ± 3.0 PSU; upwelling band, cluster II, 35.5 ± 0.2 PSU; offshore waters, cluster III, 34.2 ± 0.4 PSU; mangrove outlets, cluster IV, 32.7 ± 1.3 PSU and mangrove creeks, cluster V, 33.5 ± 0.6 PSU). The index of multivariate dispersion (IMD) illustrated high variability in zooplankton standing stock for mangrove/sea locations relative to the bay. Concurrent observations on phytoplankton revealed the importance of diatoms (r: 0.640, P ≤ 0.05). Within the eddy-generated band of upwelled water, a significant top-down control of diatoms by herbivorous zooplankton resulted in a comparative increase in abundance of dinophyceans. Based on zooplankton abundance data and species association patterns, it was possible to distinguish different zooplankton/copepod communities in accordance with mesoscale variability in physical, chemical and biological processes under tropical conditions. This was confirmed through canonical correspondence analysis (CCA) that represented coastal-offshore waters and the Bay environment in this area.     

Size-dependent, spatial and temporal genetic variation at a leucine aminopeptidase (LAP) locus among blue mussel (Mytilus galloprovincialis) populations along a salinity gradient

Biochemical genetic variation at a leucine aminopeptidase (LAP) locus is related to salinity variation in several marine bivalve molluscs. This paper details an investigation of the Long Island Sound model of LAP selection (LAP genotype-dependent mortality occurring among newly settled Mytilus edulis mussels) carried out in 1997 among three M. galloprovincialis mussel populations along the salinity gradient of Wellington Harbour, New Zealand. Significant LAP genotypic heterogeneity was observed at the LAP locus between small (<25 mm shell length) and large (>25 mm shell length) M. galloprovincialis from Petone and Eastbourne (the two sites experiencing the greatest salinity variation), whereas genotypic heterogeneity was not significantly different between small and large mussels from Seatoun (the site experiencing the least salinity variation). The Lap ³ allele decreased in frequency and the Lap ⁴ allele increased in frequency at Petone and Eastbourne, whereas the Lap ³ and Lap ⁴ allele frequencies remained effectively constant at Seatoun. Both these findings are consistent with the Long Island Sound model of selection. At all three locations, the Lap ^3,3 genotype decreased in frequency from small to large mussels, whereas the Lap ^3,4 genotype increased in frequency from small to large mussels. All other LAP genotypes occurred at low frequencies (<0.10) at all three locations and showed no evidence of frequency change from small to large-size mussels nor evidence of clinal change among the three locations. These genotype frequency data possibly indicate that the Lap ^3,3 genotype is at a selective disadvantage compared to the Lap ^3,4 genotype at all three locations, and that this selective disadvantage is related to the extent of salinity variation which exists at each location. Further investigation is required before it can be determined if the Long Island Sound model of selection best describes the size-dependent and location-specific changes in LAP allele and genotype frequencies along this salinity gradient. Comparison of the population genetic structure at the LAP locus in 1995 and in 1997 revealed a profound change from heterozygote excesses to heterozygote deficiencies for all three M. galloprovincialis populations. The reason for the change is unknown, but the change indicates that population genetic structure at the LAP locus is highly variable in time, but consistent in space, among these M.␣galloprovincialis populations. Received: 5 February 1998 / Accepted: 27 May 1998     

Food utilisation of pelagic mysids, Mysis mixta and M. relicta, during their growing season in the northern Baltic Sea

The dietary habits of the pelagic mysid Mysismixta were studied during its growing season at an open sea location in the Gulf of Finland, the northern Baltic Sea. Stomach samples were taken twice a month from June to September 1997. The most abundant phytoplankton taxa in the stomachs were diatoms and dinoflagellates, and copepods and cladocerans were the most abundant zooplankton identified. A clear change was found in the diets during the study period. Small mysids (3 to 6 mm) fed on sedimented phytoplankton in the early summer (90% benthic particles in June) but shifted gradually to a more pelagic and carnivorous diet (>40% pelagic particles, consisting of ca. 60% zooplankton in September). Seasonal changes in mysid capture ability as well as food availability were suggested to affect the diet composition of mysids during their growth. The ratio of pelagic and benthic food particles could – irrespective of the season – be explained by mysid size, whereas the zooplankton:phytoplankton ratio was better explained by season. The stomach analysis suggests that the mysids needed to attain a threshold size of 8 to 11 mm to initiate feeding on the more evasive copepods. Mysids also started to grow faster at the same time as the proportion of copepods increased in the diet, which suggests that copepods are an important energy source for M. mixta in late summer. Finally, a comparison was made between the M. mixta diet and that of the less abundant M. relicta. The diets of the two pelagic mysid species overlapped by 75% (Schoener's index). The main difference was due to M. mixta eating more zooplankton and pelagic material than M. relicta. Received: 15 September 1999 / Accepted: 18 January 2000     

Intra- and interannual variability in the larval fish assemblage off Gran Canaria (Canary Islands) over 2005–2007

The present study provides the longest and most intensive plankton and larval fish seasonal variability analysis in the Canary Islands and forms a basis for understanding life cycle scheduling and interactions among species, as well as the potential variability in transport processes of early life stages. Larval fish assemblages were studied weekly at Gran Canaria Island, Canary Islands, from January 2005 to June 2007, which represented two contrasting hydrological and biological periods. The former year was characterized by lower temperature and salinity that increased through 2006 and 2007. In contrast, chlorophyll concentration and mesozooplankton biomass decreased through the same period, especially when only the late winter bloom period was evaluated. However, ichthyoplankton abundance did not exhibit any clear pattern, as larger values were observed during 2006. The larval fish community of this oceanic island, located near the NW African upwelling, was composed of both neritic and oceanic taxa. Two families accounted for almost half of the collected larvae: Clupeidae (21.9%) and Myctophidae (20.5%). Although total larval concentration did not exhibit any seasonal peak linked to changes in zooplankton, the ichthyoplankton composition gradually changed during the year due to the high diversity and extended spawning periods of the fish species represented in samples. “Winter” and “summer” larval assemblages were identified, corresponding to the mixing and stratification periods of the water column, respectively. These assemblages were characterized by changes in the contribution of the most abundant annual taxa (Sardinella aurita, Cyclothone braueri, Ceratoscopelus spp. and Gobids) and by the presence of larvae of winter (Pagellus bogaraveo, Pomacentridae sp1) or summer spawners (Pomacentridae sp2, Trachinus draco, Arnoglossus thori, Tetraodontidae sp1). Upwelling filaments shed from the NW African coast reached the sampling area three times during this study, but changes in the local larval community were only detected in August 2005.     

Feeding and predation impact of two chaetognath species, Eukrohnia hamata and Sagitta gazellae, in the vicinity of Marion Island (southern ocean)

The predation impact of the two chaetognaths Eukrohnia hamata and Sagitta gazellae on mesozooplankton standing stock were investigated in three depth layers during two 24 h stations occupied in the vicinity of Marion Island in late austral summer (April/May) 1986. The zooplankton community at both stations was dominated by small copepods (Oithona spp., Microcalanus spp.), which accounted for >95% of total zooplankton abundance. Chaetognaths comprised <2% of total zooplankton abundance. E. hamata constituted >95% of the total chaetognath stock. The general trend in both species was decreasing abundance with increasing depth, which appeared to be correlated to the distribution of copepods (r ² = 0.45; P <0.05). Gut-content analysis showed that copepods (mainly Oithona spp., Calanus spp. and Rhincalanus gigas) and ostracods were the main prey of both species, accounting for 87 and 61% of the total number of prey in E. hamata and S.␣gazellae stomachs, respectively. In the guts of S.␣gazellae, pteropods (Limacina spp.) and chaetognaths were also well represented. The mean number of prey items (NPC) for E. hamata ranged from 0.02 to 0.06 prey individual⁻¹ which corresponds to an individual feeding rate (Fr) of between 0.05 and 0.12 prey d⁻¹. For S.␣gazellae, the NPC values were higher, varying between 0.04␣and 0.20 prey individual⁻¹, or between 0.15 and 0.76 prey d⁻¹. The daily predation impact of the two chaetognaths was estimated at between 0.3 and 1.2% of the copepod standing stock or between 7 and 16% of the daily copepod production. Predation by S. gazellae on chaetognaths accounted for up to 1.6% of the chaetognath standing stock per day. Received: 26 November 1996 / Accepted: 31 October 1997     

Epibiosis on cerith shells in a seagrass bed: correlation of shell occupant with epizoite distribution and abundance

The cerith Cerithium atratum (Born 1778) is an abundant gastropod in the seagrass beds at Cabo Frio, Brazil. In order to estimate the ecological importance of cerith shells as a rare hard substratum in the seagrass bed, the abundances of C. atratum and of cerith shells occupied by hermit crabs were quantified. The mean densities of C. atratum and hermit crabs were 1887 and 100 individuals m⁻², respectively, and these provided 0.5 m² shell area m⁻² available for epizoite colonization. The tube-forming polychaete Hydroides plateni (Kinberg 1867) and oyster Ostrea puelchana Orbigny, 1841 were the dominant visible epizoites on inhabited cerith shells. These epizoite populations were compared in order to investigate whether the temporal and spatial patterns in the epibiotic community were related to ecological and behavioral aspects of the occupant species (cerith or hermit crab). Larger cerith shells had a greater abundance of epizoites. Each epizoite showed a preference for a different occupant of the shells (the oyster for C. atratum and the polychaete for cerith shells occupied by hermit crabs). The oyster showed a seasonal pattern in abundance on C. atratum, being more common in fall (March–April). The distribution of the epizoites on the shells depended on the shell occupant species and was probably related to their different foraging activity –C. atratum ploughs half buried through the sediment, while the hermit crab crawls on the sediment surface. In both cases, the activity of the shell occupant was considered to be beneficial to the epizoites, as empty shells and shell fragments did not support a macroepifauna. Received: 1 May 2000 / Accepted: 8 August 2000